The Journal of School Education Research, 2007, Vol.19, pp.53-75 53
Predominance of Female, Slender Leaf and Formation of Adventitious Roots in Riparian Willow Populations toward Downstream Environment in the Southwest Area of Japan
' Yoshio Ishikawa (Teacher of High School at Higashi-Matsayama) Hiromichi Fujino (Teacher of Elementary School at Kanagi) Keiji Komeda (Teacher of High School at Kitayamato) Yumiko Sakaori (Teacher of Elementaiy School at Nishibefu) Osamu Yamaguchi (Environmental Sciences at Hyogo University of Teacher Education)
A total of 21691 individuals of riparian willows were examined for species distribution, species diversity, mean of leaf shape (LfW), mean number of filaments of male fiower and sex ratio. Correlation analysis was done between these ecological characters.
Rate of vegetative growth was also examined by formation of adventitious roots, using a single whole and halfofleaf. The samples are from a total of 10 main rivers of our country from Hokkaido to Shikoku. A total of 24 species and a single plausible interspecific hybrid appeared. Excluding minor or locally endemic or cultural origin species, they are classified into 3 major groups, The species in the northern group grow in the summer green vegetation zone with WI of less than 85, and those in the southern group grow in the evergreen broad-leaved vegetation zone with WI of more than 85. The third group distribute widely
in the above both vegetation zones. The species.diversity was centered in midstream regions in Hokkaido, but it has an elevational
gradient in the southem group, and the high H' values are centered in downstream regions, There is a significant correlation between H' and WI. Mean of leaf shape showed a positive correlation with WI especially in the southern area. Slender leaves of such as S. pierotii and S. gilgiana are dominating in downstream regions in the evergreen vegetation zone. Similarly, mean number of filaments has a positive correlation with WI in the evergreen vegetation zone. Systematically old Salix species such as chaenometoides and subLfragilis are also adaptive to the warmer riparian environments, because they have the high frequency of mixed flowers to be O.Oll. The mixture of female and male flowers in a single individual is able to produce fertilized seeds and this seems to be compensatory to difficulty of outcrossing in frequently flooded conditions. Predominance of females was
observed to be O.62 in fi;equency as a whole. The frequencies of females are correlated positively with WI values in the evergreen
vegetation zone. Salix species, such as S. pierotii or S. eriocarpa, is popular in the downstream regions. These sex ratios are
O.84 and 1.00, respectively. They showed a high rate of vegetative growth, and their leaves forrned adventitious roots as frequencies
of O.71 and O.64, respectively. These jointed adaptive advantages seem to the strategy of willow poulations to warrner disturbed tt fiparlan . envlronments. . ' Key Words: Riverside willow, Sex-ratio, Slender leaf, Rooting from leaf, Environmental education
Yoshio Ishikawa is a Teacher of High School at Higashi-Matsuyama, l-6-10 Matsuyama, Higashimatsuyama-Shi, Saitama 355-OO18 Japan, E-mail:[email protected] ' . Hiromichi Fujino is a Teacher of Elementary School at Kanagi, 1541-5 Shimokihara, Kanagi-cho, Hamada-shi, Shimane 697-O121,
E-mail:[email protected] Takashi Komeda is a Teacher of High School at Kitayamato, Uemachi, Ikoma-shi, Nara 630-O131 Japan . Yumiko Sakaori is a Teacher of Elementary School at Nishibefu, 543-175 Shinobe, Befu-cho, Kakogawa-shi, Hyogo 075-O131, Japan, E-mail: [email protected],ne,jp Osamu Yamaguchi is a Prof. of Environmental Sciences at Hyogo University of Teacher Education, 942-1 Shimokume, Kato-shi, Hyogo 673-1494 Japan, E-mail:[email protected] 54 The Jouimal of School Education Research, 2007, Vol.19
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JfiM aliill tiK.kgllZLRAXLLIM'eg.e}kK¥iki'Ag'ii (mpN), +355-OO18 t5il.kMM"AXLLITIiMXLIIEITI-6-IO, E-mail:[email protected]
maeq wsLts 'at?MiiiilZ*-axfjxe}:K.XE`"p:rJ, i6g7-O121 UX.LIgk'ctEE]TiiStwEfl]'-Finlpt.1541-5, E-mail:[email protected] )kne ena AR.kMgdkJf nelfiJts`ce ]r: huilfJllilf[!ZthS[JMvjNpt:r-ESXi ¥ fiErAEe, +67s-o131 #.vak9Duil!iJlliiriS[JINffIIffinyitZLs74-17s, E-mail: [email protected],jp LIJm Ir.ft..-. : ;i.lva#EJ)E ' INTRODUCTION were found in a single ear, it was described as mixed or herrnaphrodite. Sex ratio is described by a total of females . Willews grow frequently beside ponds and marshes, along among a total of female and male individuals. Mean number rivers, or even on river floodplains, especially in the cool offilaments ofmale flowers was estiniated by the weighted temperate vegetatiop zone. They are faced with frequent mean of the values of constitutive species for each ST disturbances of flood and freshet. In these habitat conditions, (Kimura 1989). For each willow species, the mean length only the restricted tree species are able to survive, such as (L) and width (W) of leaf shape excluding stipe were willows or alders. Therefore, they are called pioneer plants measured using 10 leaves. Mean leaf shape (LfW) was (Ishikawa 1980, 1982, 1988; Niiyama 1987, 1989, 1995). estimated by the weighted mean of the values of constitutive They bloom in early spring, and bear ffuits in early summer, species. Species diversity was described in three ways; in which no other plants proceed their reproductive activities. number of species (S), the evenness index (J') (Pielou They are deciduous, and they distribute popularly in cool 1966), and the general diversity index (H') (Shannon and temperate climate zone in the northern area of our country. Weaver 1949). Willows have a tendency of rapid formation However, willows grow also in the warm temperate zone of adventitious roots (Saito 1994). The frequency of this in which the potential vegetation is evergreen broad-leaved root formation was measured by using more than 30 forest. They seem to be completely dioecious (Kimura, branches. The same experiments were carried out by using 1989), but a biased sex ratio owing to few male trees, such 30 mature whole leaves and 30 half-cut leaves, respectively. as in Salix pierotii, are encountered on estuary floodplains. The experiments were taken in water containing 111000 S, eriocarpa lacks completely male individual (Kimura, diluted Hyponex solution (Hyponex Japan Co.), and in an 1989). Abnoimal sex ratios also have been observed in artificial climate chamber at 25 OC and under 10000 lux willow populations (Grant and Mitton, 1979; Crawford for 12Ll12D (Biotron LPH 200, Nippon medical & and Balfour, 1983; Takehara, 1989). There is a possibility of chemical instrument Co.). Observation was done for up to some mechanisms to compensate this selective disadvantage 30 days. The sarne experiments were done also in 6 control offew or no fertilized seed production. A total of ten main tree species of Alnus 1'aponica, [flmzis pam,ifolia, Clethra rivers have been examined for distribution of sex ratios barvinervis, Camelia J'aponica, Ilex rotunda and Eurya and mean leaf shapes and another adaptive characters in 1'aponica. All the above examinations and experiments willow¥populations. Partial hermaphroditism and extensive were done in from 1997 to 1999. ' vegetative growth trait are also reported. From these results, The warrnth and coldness indices (WI and CI) of each an adaptive strategy of riparian willows to warm temperate ST was estimated from the data reported by local climatic condition is discussed. meteorological observatories and it was adjusted with a decrement ofO.6Åé at every increment of 100m in altitude MATERIALS AND METHODS (Kira 1948). River bed gradient (RBG) was estimated by ' the maps published by Geographical survey institute of ' ' A total of 1O ' rivers were examined for willow distributions Japan, and was expressed as meters per kilometer of and its sex ratios along each main stream; Shiribetsu river corresponding river length. '' and Tokachi river in the Hol Tabte 1. Distribution of individual numbers of witEow trees and ecotogicat characters in each' station atong the nvers examined Riverandstation ShiribetsuRiver Tokachi River STIST2 ST3ST4 ST5ST6 ST7ST8ST9STIO STI1 ST12 STt ST2ST3ST4ST5ST6$T7ST8ST9STIOSTII Salicaceousspecies Ne.ef Rootin Warmthindex 625623 61.0600 58159.0 56554953.3 533 seo 44.] 457 '47549.2510527585580565566 576545 filsmant ftom Co[dnessindex -339-34.2 -34.5-373 -399-40.3 -421-44.3-4B.4 -465 -49.4 -58.3 -468 -470-47.1-473-474-428-454-479-445 -410-439 inmele leaf Riverbedradient{mlkm 10 90 25J 04 0305 2.6 fiower Meanno.offiIaments 32401.9186030 162.2 2ose4o 34 61 IS 03030510152030403060 Meaoleafshae 2322535.7 5.150 4.95.5 31222.7 5.4 5.1 5,7 6.0 lg2A212346293035 5993 Noofsecies(S) 495.65J 6 7 5 7 6.05.9605J5.0564.84.5 4343 Pielou'sindex(J') 11lt 075079 O.87082 065OJ8 081073O.73 081 O.83 O44 O79 069O.74085OS5O.91087Og8OS4'082OS2 Shannen'sindex(H') 12112755 15614766 11615367 158T42141 144 161 O72 153 1231431781962192092tl20167S1011111111196197 Choseniaathutifofia 5 o Fernalefrnalefrnixed 2f4106f4fO131291012!22fO419!O15!321081i3fO Unknown Sahhrhultenii 2 o Femalelma]elmixed 3f210 1/OIO110/O 4!3/O3fOIO i!3!O tlOfO 31210 o/efo 1101041110 Unknown 1 1 3 1 3 1 11oloolvoe!1/oo!11oolllo 27 roisusuuthanihna 7.5 e Femalelmale/mixed 5!7!O 3/6fO7f9fOOf3!OOIO!O114fO 518fO1!2/O Unknown 1 111¥302 Populusmeximowici 30 o Female!malelmixed 21voo/11o1/e!o OfllO 1/1!O ofo/ovolo2fvovolo2/oleo!e!o O/2fO1!OIO Unknewn 1 15 49 3 orrH Safiroride 2 o Femalelmale!mixed 24/291013f20/O VOIOIVIOfO 211/O11171031410 V410 5!14/O 313fO 4!6107!5/O4!6/O14A41014flllO6/5fe 1/2/OO!OfO2e59 s 22 Unknown 3220S10" tr Satimiabeana 1 O.10 Female/male/mixed 811VOlf3!O 313 313 15!7!O 713/O8!3/O34f12fO26/161015!17/O37122fO16flOfOS13fO 110fO112fO 16 e Ol g Unknown SatiJ'essoensis 2 O.55 Femajefmale/mixed 21 g Unknown oo Satlhrpetisusu 2 O.07 Female/malelmixed 17115!O94!26fO 22/15/O23!19/O 131111023120!O20!16!O34!361036!21!O 6513410 58140fO 2fllO 3013110 381311036!25/O58!26fO59136/O33112/O36/22/O22!20/e11/1!O18/10fO22112!O Unknown 11 14 46 2 o e Sahbesachaftiensis 2 O.10 Femalefmalelmixed 61154fO48/31fO 70143/O57/3410 51!331052/30/e 58140/O52f39/O351341031!2710 21/2010 4V3610 63!32/o t05f60!154!211e34/331038A5/O29/121252128fO41f16102e/11/O16f17!e2S126/O Unknown 34 42 22 3 17 22 o e6 g Saliihtegra 1 O.02 Female/male/mixed 111fO10!4!O 29/20!O56138!e117 64/46/19f7fO 39/241016f15/O13/11!O 3712511 33f15/O 1313/O 1417/O7!1/O20!9!e2111810261151023/12!e31123!O9/10fO 712/O7/710 172 ifg 1 4 o 02 Unknown 74 Saligraoifisty/e 1 O.10 Femalefmarelmixed 13AOIO 1112fO3,fO!O12f2101219!O12!21014141047!19!043f20!O 18!2SIO12/11fO 21 o grt¥ Unknown oo Salibesubfragr7ib 3 O.33 Femalefmale!mixed 49150f433151/1 161t2fO4f1410 3171011!22fe 4f19fO1!1/O513fO 4/5!2 5151126114!e7120109131!111!28fl10!14fO4!4fO3!3fe efve1!QfQ' Unknown 2 e oo Safiiserissaefolia 2 o FerrSalefmalelmixeci 210 oo f Unknown Salipierott;' 2 O.71 Femelelmalelmixed S'g Unknown Salixerioearpa 2 e.64 Female!mate!mtxed Unknown tY. SalibegiYgina 1 O.11 Femelelmalelmixed Unknown Salicheenomefoide 4 O.05 Femaiefmalelmixed Unknown 't Salixyoshinoi 2 O.02 Femalefmale!mixed Unknown Saltkbakko 2 o Femalefmale/mixed Unknown Selikm'uyanagi' 2 O,08 Female/Fnale!mixed Unknown Sefiisiebotdiana 1.5 O.06 Female/male!mixed Unknown SabbekoriYanagi 1 O.07 Femalelmale/mixed Unknown Seltivbaby/ont'ca 2 o.le Female/malefmixed Unknown Safixmatsudana 2 e Female!male/mixed Unknown Hybrid1 1 Femele!male!mixed Unknown Totat Female l3e186 164153 133107 12911793 135 124 49 l38 18D135169178145200i89ID5 82S5 MaJe 131115 121125 94 97 42 8S 10274 M[xed 9789 1049973 1 o o 1086410814010714912667 Unknewn 47 136 30 1126 2413 357 t21 229 3390oo Total 28232841 294313935oo 25520910 249233i7316177 277 29019928032t275357336IS5 217249 Sexratio 051062 058055 058O.55 055O.54056 O59 O.56 O54 061 O.63O.68061O.56O.58057O.6061 O.45O.54 Tab{e 1. Continued 1 Riverandstation TekachiRiver ShinanoRiver STI2ST13ST14 ST15 STIST2ST3ST4$T5ST6ST7ST8 ST9STIO STII ST12 ST13 ST14 STI5 ST16 ST17STI8 ST19 Salicaceou$species No. of Reetln Warmthindex 468 1069101.3101.2100.81003999995 974 953931 878 825 833 841 892 906 941 filsmant 524514504 -77.6 920933 ffom Co]dnessindex -54,4-56.9-¥62.7 -- 5.3-g.S-8.7-9.1-9.6-10.0-104 -12.1 -138-154 -180 -205 -203 -20O -207 -195 -183-17t -158 in male Ieaf Riverbedradient(m!km 1203.070 231 21 57 40 77 16 12 09 os fiowet Meanno.offilaments 70 02O.202Oi030607 18 2438 17 14 IS 17 18 18 1109 25 Meanleafshae 503.060 4.8 1718202i201617 61 1617 58 47 47 52 62 57 2421 49 No.ofsecies(S) 525.746 4 615.85956546362 8 6457 7 7 8 8 8 6 5355 6 Pie(ou'sindexJ'} 052065087 O70 074077067066073060055O78 073086 O89 O84 OS2 085 080 O90 089073 O57 Shannon'$index(H') 094i26169 097 163150130128142118107161 14215476 174 163 17e 176 i66 162 123iOt44 102 Choseniaarbutifolia 5 o Female!male!mixeci Unknown Salixhulteni)' 2 o Femalelmalefmixed 2f3/O5fllO 3fOIO i Unknown 8, Tot'susutirbeniann 7.5 o Femalefmalelmixed 10f610 OIOIO Unknown oo 68 l5 o Popu/usmaximoruicz 30 o FemalelmaLefmixed o!ofovofo21o!e e/o!o e Unknown 15 P.. po= Satirorida 2 o Femalelmale/mixed 126191017!O = Unknown 2 SatimtYabeana 1 eno Femalefmalefmixed g Unknown SatrbeJ'essoenstS 2 O.55 Femalelmaie!mixed 26110/'O 17!t21020/2VO 712210 611310 4/1VO 71910 tlOIO Of210 Unknown 4A!OOIYOOA!O2/O/Oo o o o o o o g Sal/hrpet-susu 2 o.o Female!male/mixed 3311VO3211410 vo/o4/2/o2/ofe O!1/O Unknown oo o g Setisachefinensis 2 O.10 Female!male/mixed 52!30fO651341043!20!O 2012310 310/O 211!Oi211510 1514!O 101710 6felo lsfive 3!110 O1210 Unknown e6 24 o o o o o o o 2- 1 VOIO2!OIOV070 Salixintegra O.02 Female!male/mixed 212106f9!O1!3!O 3fO!O2fOfOOII!O110!O VOfl 3/O!O6!710 9flllO 12f7fl 21!14!O 7!2!O 110!O 5!1!e IAIO OOi2 o o o o o o Unknewn oo e e Sattbegracitistyla 1 e.lo Female/malelmixed 1!O!O2!O/O671fO 2!3fe 21VO25112/O 7fOIO 8fS!O 4/efo g Unknown 2fO!OO1270011!O eoo o o o o o Sattisubfragt'lis 3 O.33 Femalefmale/mixed 611103/1!O 71101010A5!Oi2!1710181201011118fO121131021119109f1510 31410 lf2fO ef4/o O!4!O lf3/O 21610 22f12fO18127/e 3313110 s Unknown o o o o o o o o Safihrserissaefofia 2 o Female!male!mixed 8!10fO 2112103f3!1 16113!O 31710 6f8!O 3f510 9!14fl 915fO lef3/O41170 414!O I4!2fOef3!O715fO41210113106/3/O4/410 s-. Unknown oo o o o o o o o oo o Satipierotl)' 2 O.71 Femalelmalelmixed 10115!O1615101!2/O2f51Dll/12!O41611O12!O 4/3!O 3!O!O 2fllO 815!O 16/4/O 8!2!O3fVO 210!O 6'" Unknown o oo o o e o oo e g¥ Sefixeriacarpe 2 O.64 Female!male/mixed Unknewn oo Satigilgiane i O.11 Female!malelmixeci 22121!O23!i3fO19112fO16!14fO14/8fO46129!O3?134/e31!17fO 34f13/O"f8fO 719fO O1210 1/1!O O!1/O 7f210 6f6fO 4!61021!1410 611010 t-5 Unknown o o o o o o o o Salicheenome/oide 4 O.05 Femaiefmalefmixed o!yo o oo oo g Unknown Sefiiyosh,hbl 2 O.02 Femajefma]e/rnjMed eo Unknown Satibakko 2 o Femalefmalefmlxed -g¥ Unknown Salikihuyanagi 2 e.os Female!male/mixed ca Unknown Sa/ibesiebotdihna 1.5 O.06 Female/malefmixed Unknown Satikorienagi 1 e.e7 Female!malelmixed 1!'Of'OV'O!'O Unknown Sa/ihrbabylonica 2 one Femalelmaie!mixed O!lfO Unknewn o Satibematsudana 2 o Female!malelmixed O!1/O Unknown O!1!OlfO!O o Hybrid1 1 Femaie!malelmixed Unknown Total Female 63115104 23 S2 57 56 48 45 34 38 46 Male 23 62574142387066 56 6154 62 49 38 41 25 24 4446 48 1 4363 1 1 2343 Mixed 365651 o 50383742445359 o o o o o Unknown 55 e o e o e o o o Total 126107 101 139 104118Ol ll9 106 86 86 60 62 oo 94 111177262 O.59 O.4S O.53 O.56 O.52 O.58 O.61 6789 O.49 Sexratlo 064067067 05 055060e53050e46O.5705311295788482124125 059O.46oo O.66O.52oe "th th oo Tabte 1. Continued 2 Riverand$tation Shinano River MaruyamaRiver YataRiver ST20 ST21 ST.22 ST23 ST24 ST25 ST26 ST27 ST28 ST29 ST30 STI ST2ST3 ST4 ST5 ST6 ST7 STI ST2 ST3 ST4 Salicaceousspec]es No.ef Reoting Warmthlnclex 951 955 96O 719 17.1 823 753 683 613¥ 545 41.5 "e7 10991091 t081 1052 le3s 995 1"6 1056 953 883 '- -24.0 -22.3 fiIamant from Coldnessindex 156 -153 -15e -205 -244 -283 -322 -36O -399 -44 -4.7-50 -54 -63 -69 -¥ 78 -2O -62 -9S -123 inmale feaf Riverbedadient{mlkm 15 33 58 80 S5 85 116 22O 159 174 848 04 25 36 59 133 40 60 130 369 fiewer Meannooffilaments 23 16 16 20 23 19 16 18 i3 14 t9 16 Ol15 27 23 22 13 15 13 11 11 Meanieafshae 51 6.3 6.5 5J 4,7 53 51 54 5.6 4.5 54 6.0 1623 46 41 4.3 4.3 43 4.0 38 38 No.ofsecies Riverandstation MukoRiver KakoRiver IboRiver STI ST2 ST3ST4 ST5 ST6 ST7 STI ST2 ST3 ST4 ST5 ST6ST7 STS ST9 STI ST2 ST3 ST4 ST5 ST6 ST7 Saiicaceousspecies Ne.ef Reotimg Warmthindex 11S.9 114.9 110.e108.3 107.3 106.0 100.3 119.8 118.3 117.7 116,5 112.5 114.0117.4 110.8 1039 119.3 116.5 113J 1088 1037 93g 89O fllamant -3.4 -4.3 -3.1 frem Coldnessindex -¥ 5,4-6,O -6,5 -6.4 -7J -3,5 -3,6 -3.8 -3.8 -4.5-5,3 -5,3 -6,8 -3.3 -3.9 -4,6 -5,7 -6,5 -9.2 -10.3 inmale leaf Riverbedradient(mfkm 2,O 5,O 6.02.4 1.0 1,6 7.0 O,9 1.9 O.8 1,5 2,O 1.01,O 3.0 S.5 1.5 3,5 3,5 6.0 9.0 9.5 120 fiower Meanno,offilaments 1.3 1.2 1.22.4 1.7 1,8 1.4 2.2 2.4 2.0 2,2 2.0 1.92.0 1.7 1.2 2.4 2.0 1.5 1.3 10 10 16 Meanleafshae 6.7 4.0 3.93,9 4,O 4.3 3.7 4.8 4.7 5,3 5,1 5.0 4J4.6 5.4 3,7 4.9 5.1 4.5 4.1 3,6 3,7 2.4 No.ofsecies(S) 6 6 6 4 7 8 9 8 7 11 9 2 7 5 9 7 5 3 3 Pielou'sindex(J') O.59 O.47 O.40OJ2 O.73 O,54 O.51 O.59 O.73 O.79 oss 077 076087 O65 O47 O84 OS7 O66 O53 O03 O12 O43 Shannon'sindex(H') 1,05 O,83 O.721.6069 1.30 O.76 O.99 1,22 1,61 1.64 1.70 1.90 1.671,808S 1.44 O.33 1.64 1.40 1.45 1.03 O,Ol Q.13 O.50 Ohosenlaarbutifolia 5 o Femalelmale/mixed Unknown Sehbehuttenil 2 o Female!malelmixed Unknown v ToisusuurbanJbna 7,5 o Femalelmale!mixed o Unknown oa Populusmeximovvt'ci 30 o Femalelmalelnn[xed e Unknown s' Satixrori'de 2 o Femalefmalefmixed g Unknown 8 SetibemiYabeana 1 O,10 Female!male!mixed Unknown a Saltky'essoensis 2 O.55 Femalelmale!mixed tho Unknown E Salibepet-susu 2 O.07 Femajelmale/miKed N Unknown o pt Saltksachalihensis 2 O.10 Female/malelmixed O!8/O 3/410 OA!O Oll/O O12!OO!2fO lfOIO OIO/O OfOIO = Unknown o 1 o o o 3 7 pt Sefihrihtegra 1 O.02 Female!malelmixed O!110 11010 slelo 2!21022/610 O1410 m rc Unknown o o o oo o s pt SelihrgraciYi'styla 1 O,10 Female/male!mixed 42120fOI09f48fO 45!15fO 75723fO 1611910 5S/101!O O!1!O 30f9!O 27111!O 7!910 o!o/o 31flOIO37f12!O 36!20!O 4/5!O OIOIO ofo/o o/ofo O/OIO O!OIO o!ofo 9 Unknown o o o o o o o o 56 oo o o 3 IS 69 64 900 154 Seltbesuhfregtzas 3 O.33 Femalefmale/mixed 39!4010 4110!O 8!3fO O12fO Oll!O 1/OfO 2f2fO 17132/O 5f18fO28!31fO14O 6!4/O OllfO o!o!o ur Unknown ooo o o o o o o o o o 1 eq SafrbesertSsaefotia 2 o Female/maie!mixed 9!5fO Ofl!O O!VO4!4/O 23!4!O 102!O!O 101/110 52fO!O 91A/O 3610fO5410!O 6816!O OfOIO OIOfO OIOfO OIOIO OIOfO B' Unknown o o o o o o o oo o 12 32 9 2 2 SatipJ'erotl)' 2 O,71 Female!malelmixed 911tlO 210!O 6!OfO4!OIO 2/O!O 101010 t82!410 136/110 46/O!O 1710/O 1!110 11VO OIOIO o!o!o o/o!o OIOIO :. Unknown o i oe o o o o o o o ooo 51 7 11 le vee Salibeertbcarpa 2 O.64 Femalelmale/rnixed 22f070 s/o!o 34fOfO35fO/O 531010 105fO!O 27!O!O 5910fO 47!O/O 7910fO 41!O!O 72!OIO 41fOIO43fO!O 38!OIO ofo!o O!OIO ofofo O!OIO OIOIO as. Unknown o o oo o o o o o o o o o 27 43 31 6 1 pt Saltbegilgiana 1 O.11 Femalelmalefmixed 68!7510 O!110 210/O 41010 Of310 Of510 o!vo OIO/O o!o/o o!o/o Q Unknown o o ooo o o ooo o 3 2 2 SalZchaenomelolde 4 e.os Femaie/malelmixed O12!O lfOIO2!8!O 312!O 110/O 314fD 161101e 15121fO 4f410 613fO 3f3fO O12!O3!610 O/OIO OfOIO OfOIO o!o/o -o Unknown o 10 4 20 15 7S 19 19 21 32 8 3 1 Salibeyoshihoi 2 O,02 Female!maielmixed 4/2/O 4!lfO 2fO!O1!O!O123 112!O 1!O!O 2!OfO 110/O o!o!o o!o/o OIOIO vo Unknown o o 4 o o 6S o 7 3 4 v Sati)cbeRko 2 o = Female!ma:e!mixed O!OIO pe Unknown oo 1 o Sahbekthuyanagi 2 O,08 Femalelmalefmixed o/o!o s Unknown 2 Sattbesi'eboldlane 1.5 O.06 Female!malelmixed OIO!O Unknown 47 Satibekorivenagi 1 e,o7 Femaie/male/mixed Unknown Sehbebabytonice 2 O.10 Femaie!malelmixed 11010 Of310 o/o!o OIO!O o/o!o Unknown o o 9 2 1 Sahbematsudana 2 o Femalelmale!mixed Unknown Hybrid1 t Female!malefrnixed Of410 Of9fO Ol1210 O/8fO O12412 Unknown o o o o o TotaÅ} Female 112 56 152i32 139 124 210 281 331 258 125 191 117187 150 4 o o o o o o o Male 95 23 43 19 t14 25 42 29 22 64 38 6 o o o o o o o Mixed o o 4968 o o o o o o o 2 3562 o o o o o o o o o Unknown o 1 10 4 25 15 78 t9 19 77 o o 141 108 133 89 906 158 55 Total 207 80 202223oo123 192 147 349 321 451 306 166 334 172257208oo 188 ie 141 1OS 133 89 906 158 55 Sexratio O54 071 076066 O76 O87 065 092 O89 O90 OS5 O74 O.68076 O.80 O.40 eth oa Tabte t. Continued 4 Riverandstation KinoRiver Shimanto Rlver STI ST2 ST3 ST4 ST5 ST6 ST7 ST8 ST9 STIO STI1 ST12 STI ST2 ST3 ST4 ST5 ST6 ST7 ST8 ST9 $TIO STII Sa]icaceeusspecles Neof Reeting Warmthjndex 130.7 1340 127.6 125.5 123.1 114,9 116.5 114.0 103,6 97.2 86,5 80.1 127.0 129.0 125.9 124.S i24.6 120.2 122.1 120.S 119.5 117.0 115J filamant frem Coldnessindex o.e o.o -1.0 -1.7 -2.6 -3,O -2.g -3.3 -5.1 -6.2 -a.a -9.1 o,e o,o -O.3 -O,4 -g.4 -o.s -e.s -O.9 -1,1 -1,4 -1.6 immale leaf Riverbedadient(m/km e6 11 18 20 22 21 41 24 9S 60 IOO 293 02 08 10 IB 02 20 20 20 20 40 20 fiewer Meannooffilaments 26 28 29 29 25 24 18 15 11 10 12 10 19 28 24 20 13 13 15 11 13 16 14 Meanleafshae 4.5 40 3.9 40 4.4 41 44 36 37 36 42 36 36 35 36 39 40 36 36 37 35 35 35 No.ofsecies(S) 6 7 s 8 9 10 9 s 4 3 4 1 4 7 5 5 4 4 4 4 3 4 3 Pielou'sindex(J') O.91 OJ8 O.74 OS3 O.83 075 O.71 O64 O.52 O14 O57 ooo O82 O70 081 O85 051 O37 O52 O26 O40 051 O46 Shannon'sindex(H') 164 152 154 172 180 172 t56 134 072 O20 079 ooo 113 136 131 137 e71 O50 O72 O36 O44 O70 051 Choseniaarbutifolia 5 o Fema]elmale!mixed Unknown Sefi[hultenii 2 o Fema[elmalelmixed Unknown Toisusuurbaninna 7.5 o Femalefmalelmixed Unknown Populusmaxioveici 30 o Femalefmalefmixed Unknown " Salirorida 2 e Female/male/mixed 2 Unknown Sahbemiubeene 1 O.10 Femalefmale!mixed o Unknown 5 SafiiJ'essoensis 2 O,55 Femalefmale!mixed su Unknown o. Sazapet-susu 2 O,07 Femaie!malelmixed Unknown ge Safixsecha/inensis 2 O.10 Femalelmalelmixed Unknown & Se/riihtegra t O.02 Female/male!mixed 1/efo 41010 441i7fO 14fS!O 21010 Unknown o o o o o g Sa/igreciliStyla 1 O.10 Femalelmale!mixed VOIO 7!810 9/2!O 191Sll 61A8!O 6212010102!2010 9412110 5913410 61130!e 53142!O 42118!1 1211410 B1610 34A4105112310e44f5510 66130fOI 68188!OI 6016SIO 6V3110l OS/5410 g Unknown o o o o o o o o o o o o o o o o o o o o o o g¥ Salixsubfiagr7s's 3 O.33 Femalelmnlelmixed 9f19!G 10f2SfO 4flofe 2V24fe 24!34fG 4115fO 11133fO 7123fO D/4fe 5f4fO Of2fO Unknewn o o e o o o o o o o e R Seltbeserissaefolie 2 o Femalelmalelmixed 21151e 413!e 13!2310 13A510 17/22!1 7f3310 O/2/O 112fO Unknown o o o o o o o o g Selipferotir' 2 O.71 Femalelmale/mixed 14/710 i51010 161310 251510 913!O 9!510 61410 O!210 7/110 8/7f2 12A110 S14/O 1211110 19!3!O 5101e 7/OIO 201110 71e!o s!e/o 7fO!O Unknown o o o o o o o o o o o o o o o o o o o o 'z Saliertocarpa 2 O.64 Femalelmale/mixed 41!OfO 41/O!O 24!O/O 28/elO 23/'OIO 13/O/O 51010 1/OfO 2/OfO vefo Unknown o o o o o e o o o o .- Sefigttgtbna 1 O.11 Female!malelmixed 21110 Oll/O 1!OfO OlllO V2fO 2011110 210/O 7/1210 , Unknown o e o o o o o o 4 O.05 10/18/e 4915210 4214910 16124fO 31/47!O 3!8!O 9/121e 1!210 711316 41/38/4 13/13!O 110fO 811310 6/712 16/1OIO 8f1210 8/14!O Sahbechaenomeloide Female/male!mlxed 21!1810 ofve 5AlfO 2A/O tL Unknown e o o o o o o o o e o o o o o o e o o o o Se/ixyoshihoi 2 O.02 Female/malelmlxed OlllO 2!210 313!O 2/410 olve 41310 1711410 12/11/O 3!OIO 5fO!O 4fOIO 1/1/O 11110 Unknown o o o o o o o o o o o o o v Selibekko 2 o Femalelmalefmixed Unknown Sahhrkihuynnegi 2 O.08 Femele/malelmixed ef4!o e/21o O1210 Unknown o e o Sa/tlsi'ebofdiens 1.5 O.06 Femalelmalefmixed Unknown Safukoriunegi 1 O.07 Femalefmale!mixed 210fO Unknown o Sahbebabyfonice 2 O.10 Female/malelmixed Unknown Safimetsudane 2 o Femalelmalelmixed 1!O/O Unknewn o Hybridl 1 Femalelmalefmixed Unknewn Totai Female 78 92 114 140 "2 129 117 165 115 61 74 53 61 85 23 72 74 162 83 l91 183 75 123 MaLe 6e 51 97 101 95 126 79 78 40 35 4B 42 41 79 21 49 2G 68 37 91 IS 44 6S Mixed o o e o 2 o o o o o o o 9 4 o o o o 2 o o o o Unknown o o o o o o o o o o o o o o o o o o D o o o o Totai 138 143 211 241 209 255 196 243 I55 96 122 95 11i 16S 44 121 1OO 230 122 282 26t 119 19t Sexratio O.57 O.64 O.54 O.58 O,54 O.51 O.60 O.6S O.74 e.64 O.61 O.56 O.60 O,52 O,52 e.6o O.74 O.70 O.69 O.68 o,7e O,63 O.64 Tabte 1. Continued 5 Riverandstation ShimantoRiver Total Sexratio Chi-square STt2ST13STi4ST15ST16ST17STIB$T19 Salicaceousspecies Ne.ef Rooting Warmthlndex 114.4114.6112.5110.6109.3105.5iO0496.6 filament from Coldnessindex inmale leaf Riverbedradient(m/km -18-1.8-20-23-24-29-36-41 flower Meanno.offilaments 202.0103.02060BO300 Meanleafshae 13101010101010i2 No.ofsecies(S) 363.6363.636363629 Piete"'sindex(J') o46ooooooeoooooooooooogs Shannon'sindex(H') oslooooooooooooooooooo6e Choseniaathutifolila 5 e Femalelmalefmixed 6e!113!O e.35 16.24 Unknown 14 *** Sahhrhu/tenii 2 o Femalefmalelmixed 33/19fO O.64 O.85 Unknown 21 NS w Tot'susuurbeniana 7.5 o Femalefmalelmixed 32/45fO O.42 2,21 8 Unknown 110 NS g Populusrnaximorvicz 30 o Fema[e/malelmixed 1417!e O.67 2.38 e Unknown 267 NS s' ge Sahhrrorida 2 o Femalelmale!mixed 134114810 O.48 O,72 as Unknown 74 NS o SalixmiYabeana 1 O.10 Femalelmalelmixed i771109fO O.62 16.17 o Unknown 3 *** th Salibefessoensib 2 e,55 Femate/malefmixed 94/109/O O.46 1.li oth Unknown o NS e Satipet-susu 2 O.07 Femalelmale!mixed 888!537/O O.62 86.46 su o Unknown 102 *** po Salisachaiinensis 2 O.10 Femalefmalefmixed 1411/93713 O.60 95.69 = Unknown 139 *** pt Salibeintegre 1 O.02 Femalelmalelmixed 826f482/4 e.63 90,D6 m- Unknewn 8 *** o: pt Satlbegraciiistyta 1 O.10 Femalelmale/mixed 99f501e571451090!8VO67f44/O56/531066f581063/41/O5!6fO 3237A698!7 O,66 472.66 9 Unknown 1279 *** Salixsubfragi7i's 3 e.33 Femalelmalelmixed ID23!1451fli O.41 70.34 ar eg Unknown 15 *** Salixserisaefolia 2 e Femalelrna[efmixed 9231253!3 O.79 381.72 s' Unknown 57 *** :, Sattipierotii 2 O.71 Femalelmale!mlxed ie/o!o 778/149/3 O.84 426.80 g Unknown o 80 *** Satlbeeriocatpa 2 O.64 Female!malelmixed 93VO!O 1.00 931,Oe g' Unknown 108 *** g¥ SaligiYgiana 1 o." Femalelmaie/mixed 677/532fe O,56 19]6 Unknown 7 *** O- Sabbechaenemetoide 4 O.05 Female/malelmixed 7/810 382f466f12 O.45 8.32 s o 268 Unknown ** g¥ Sa/tiyoshihof 2 O.02 Femalelmale/mixed 84/4810 O.64 9.S3 Unknewn 18 ** :- Salibakko 2 o Femalelmale!mixed 6/110 O.86 3Jl g¥ Unknown 1 NS Sa/ixkiuyanegi 2 O.08 Femalefmalelmixed Of8!O o,oo 8.13 : Unknown 2 ** Salibesiebo/dilana 1.5 O.06 Female/malefmixed 315!O 4!5/e O.44 O.22 Unknown o 47 NS Safiikorilenagi 1 o.e7 Fema[efmaLelmixed 710!O t,oo 7.t4 Unknown o ** Safixbebytonica 2 O.10 Femalefmalefmixed 3f5!O O.34 O.63 Unknown 12 NS Saiimatsudena 2 o Female!male/mixed 5!210 Oll 1.43 Unknown o NS Hybrid1 1 Femaielmaielmixed OA60!3 o.oo 160.el Unknown o ** Total Female 11729 Male "65790675666638 7284 Mixed 5845Sl44535841" 46 Unknewn 2632 Total 21691 O62 1038.67 174t0217111110912410419 O.62 Sexratio O.61e.56O.53060051OS3061O.42 *** a- 62 The Journal of School Education Research, 2007, Vol.19 RESULTS to certain locality and cultivated origin, the above 13 popular species were examined in 'the distribution of their Characteristics of distribution of riparian willow sex ratios and RBGs. Most of them were seen in STs with populations along river low RBGs or floodplain in another word (Fig. 1). The A distribution of willow populations was surveyed in four wide distributed species have a wider range of RBGs. atotal of122 STs along 10 major rivers from Hokl(aido to There exists also a wide range in sex ratio among species. Kochi. A total of 21691 individual willows were examined The predominance of female is seen typically in S. pierotii for gender and leaf shapes as a result ecological adaptation to or S. serrisaefolia or another species. The ecological and variable riparian environments from downstream to upstream. evolutionary' bases df sex rario disorders are analyzed in A total of 24 willow species and a plausible interspecific the following sections. hybrid appeared. They are arranged in a sequence from Distribution of species diversity a}ong a main river northern species to southern species. The examined rivers stream and STs are also arranged in the same way (Table 1). Distribution of species diversity along each river was The indices of WI, CI and RBG are described also in the examined by 3 indices of S, J' and H'. There was no basic table. 'The WI of STII of Shiribetsu river is 44.7, and is difference between these three quantities (Tables I and quite close to 45. The other WIs of the STs of rivers of 2). Thus, the following analyses were done by using only Shiribetsu and Tokachi and the upstream regions of H' values. The maximum values of H' were seen in the Shinano river are in the range of 45 to 85 in WI. Thus, midstream regions of the river$ of Shiribetsu, Tokachi, these regions are in the zone of summer green forest as Shinano, Muko, Kako, and Kino. Those were seen in the potential vegetation (Kira 1948). In the stations from downstream regions of the rivers of Maruyama, Yata, Ibo, STIOto ST22 ofShinano river, all the WIs are over 85, but and Shimanto (Fig. 2), The former rivers are located the CIs are under -15. The corresponding potential vegetation rather in the northern and central area of our country. The seems to be warm temperate deciduous broad-leaved forest. latter rivers are in the southern area of our country. A In all of the remaining regions, the WIs are over the value high value of species diversity is indicative of high adapt- of 85, and the CIs are over the -15. Thus, they are in the zone ability to that ecological' condition. The maximum value of of evergreen broad-leaved forest as potential vegetation. 2.2 was observed in Tokachi river, and every H's in From the viewpoint of WI and CI, the appeared willows. can Shimanto river were less than 1.5. 0n the average, the be summarized in the following groups. The northern or northern rivers have higher H's than those of the southern typical species in the summer green forest zone are C. rivers. The consistent results were already reported ' arbutifolia, S. hultenii, T. arbaniana, P. maximowiczi, S. (Ishikawa ' 1980, l982, 1983, 1987; Niiyama 1987, 1989, rorida, S. miyabeana, and S. pet-susu. The intermediate or 1990). It is noteworthy that deciduous willows inhabit and possible species in the warm temperate deciduous broad- are well adapted to high WI region more than 130, where leaved forest zone is S. ]'essoensis. The wide distributed evergreen broad-leaved forest is expected as potential species are S. gracyllistyla, S. subfragilis, S. integra and vegetatlon. S.sachalinensis. The southern or typical species in the Distribution of means of leaf shapes along a main evergreen broad-leaved forest zone are S. serissaefolia, S. river sitream pierotii, S. eriocarpa, S. gilgiana, S. chaenomeloides, and S. Willows have two main types of leaf shape. Willow yoshinoi. S. bakko and S. sieboldiana seems to be immigrants species, such as, T. arbaniana or P. Tnctximowiczi has a deviating from mountainous habitat. The remaining 4 species, typical round-shaped leaf, and S. serissaefolia or S, gilgiana S. kinuyanagi, Sl koriyanagi, S. babylonica and S. matsudana, has a typical slender-shaped leaf. A distribution of mean seem to be also deviates from cultivated fields. One plausible leaf shapes was examined along main river streams. All of the hybrid willow species was observed in Kako River. The rivers showed the identical direction of the predominance male flower has ohe or two filaments like S. sieboldiana, but of slender-shaped leaves at downstream regions and that of it has slender leaves like the intermediate between S. pierotii round-shaped leaves at upstream regions (Fig. 3). No and S. serissaefolia. There exists completely no female morphological differences were observed in the leaves of flower of this hybrid. The plausible hybrid willow was male and female plants. The consistent results are reported tentatively named Hybrid 1 in this report. in arctic willows (Craford and Balfour 1983). After exoluding the willow species seen rarely, endemic Predominance of female and slender leaf in riparian willow populations ' Distribution" of means of numbers of filaments in and contains the rest of the species. The species of the male flower qlong a main river stream first class, especially the first two, are popularly seen in Male flowers of willows have a variety of numbers of wide distribution, especially in the downstream regions. filaments or stamens. P. maximowiczi has about 30 filamets Their branches are fragile and easily broken by human (actually from 20 to 40), and S, gracyllistyla or S. gilgiana hands. The willows of the second class are popularly seen has a single filament. Some intermediate numbers are among in the upstream regions ofthe northern part or high altitude willow species. The n!tmbers of filaments are indicative of of our country, when the last two species are excluded. their systematic relationship (Kimura 1989). A distribution Their branches are elastic and not easily broken by hands. of the numbers of filaments per single male flower was The species of the third class are commonly seen in the examined along main river streams. Both rivers in Hokkaido intermediate regions between the above two classes. Further showed the tendency of decrement of this quantity toward experiment was done in the same way by using the half-cut downstream regions (Fig. 4). On the contrary, the rivers leaves. Basically, the parallel results were gained but less in the southern area' of our country, rivers ofKako, Ibo, Kino pronounced (data not shown). and Simanto, showed the opposite direction of increment Relationships between ecological conditions, system- o¥f the quantity toward downstream regions. The rest of the atic characfers and species diyersities rivers showed somehow the intermediate between the Some of the above independent distributions of the above tendencies or almost uniform distribution. characters seem to be parallel or to be contradictory in Distribution of sex ratios along a main river stream riparian willow populations along rivers. Correlation analysis A predominance of female was found to be O.62 as the was done among the main characters appeared in Table 1. jointed frequency of. sex ratio of all the willow species There exits a significant positive correlation between sex and all STs in Table 1(xZFi=1038.67, P $elix mlytim Saliiissoeasls Tdisasuutweh- Salixrotidd Sal1ti P"t-esV 1 1 ..e t 1 .9 t e e ,9 ! eg E og tTXO,9 fie.g 1 o,e . ts ;ca oe - o.e S o.e q i oe aog o.) Ol O.1 O.1 O.1 O,6 O.6 o.fi . afi O.6 05 el 05 os o,s e4 e.- O.4 . o.4 O,4 03 oa O,3 e.3 . 03 O.2 02 e2 e.2 O,2 O,1 o,t oi e.1 Q.t o o o o o e lo 2o 3o 4o so fio 7o en ge o 10 tc gi..,6gdfu'ien?PmxA9 a 10 e 10 20 o IO 20 30 -O SD SO 70 oo 90 R;verbedItedient(mArtt) 3o405 2o3ouasese?oeegeRivetb-dgr"dierttCmXkm) coqesofioloeegoR;vgrbedptrflent(mlkul F;vertedsr.dl.nt("Vlawo) g o tr s Sel;:eutvftte"is Selixgresilistylt Selix intest- Sali#seeFuliptas;s ¥g ' i -g ' ol g e o.g ; o.g eog . 'gD9 S ee ,-} tt os los :t Qe e to . . . e.7 . ut O.7 . O.1 oeel . 06 o" :ss o.s g 05 ¥ e.s g: "" O.4 . :l O.4 oe O.3 :; l 02 :1 O,2 :, ol "1 "g o .o o o ie 2e ao #o so fie 7o ee se se 4o so fio 7e eo co o to ao 3o 4o so ss 7e co so e lo 2o 3o co so eo 7o co go R;vet bed ereetsnt(",Ak"D e te 2o R;ver bed ItedSent (mXk"" . River bed sreeientCthiflaT" River bed sttditnt(mlk"" g -ee.tr i'9 Selicehene"mb;des SetixtilarcLil SttltsefhoeefoIta SelLx l"tist 1 .R ¥e- t ..e..1 e e 1 e e.g t09 'f og g goeut07e6os04 " oe l os S o.e va m a7 07 o? o.fi O,G os ¥ e5 0.5 . O.5 o- od . o.A U" e3 D.S 03 . e3 el O.2 O,7 O,2 O,1 o.t o.t O,1 e e o o' o O tO ' 20 3D co so 6o 7e co go e le 2o 3e 4o se 6o 7o ed ge o ie 2o 3o "o so eo lo eo go lo2D3o4osose7oeegoAverbedgrediept(mllep) Riv.tbedlr.dilant(mAkst) Aver b.d sredle"tt"Vken) R;ver bed gredieal(mAvt{) Fig. 1. Distribution profites of sex ratios of common riparian wiHow species against warmth index (WI) there in. The upper 5 specles grow ln the summer green vegetation zone with Wl values of tess than 85, The bottom 4 species grow in the evergreen broad-Ieaved vegetation zone with Wls of more than 85, and the centrat 4 species distribute widety in the both vegetation zones. Lines in each figure are the most fitted Linear regression lines. ' ' ' ' ' ' ' ' T. ttCi N th E 9. uaE g- ua g. g2g Shannonindex(H) Shannon index(H) Shannonindex(H) Shannonindex(H) Shannonindex(H) RB s¥ proa"an NNen ' N m u" = e m .- en r" ta 9¥ u e o o P.O-.i!NN o o roa Ir.o o.. oaoaoen oen.anNa¥ oa" g,:¥g ¥ s en ¥ N " ee e rtta mo - ¥ -h-m a or 6ctob" ;' n ¥ en Q ges' ¥ " ua = -. e HV O tn ttJ g o 8g2 s ¥ m s' ptgoA 7= 6 . o- g SE'l e 8 oo iggg-- g "geg'. es. ljgect-gs x .K e eto tho :lg gl¥ ff- l gl¥rt ca getr.gG e s o *.g¥ i." ct/. = ou Å} 9 6 .s' =E =oQ" " B mT 'v pt g'ua e o g y pa'- w pt tt¥ H o- cr=o B pa u. pt th ..-OM s' R di Shannonindex(H) ShannQn index(H) Shannon index(H) Shannon index(H) shannon index(H) :. :¥¥ vee 2 i¥ 9 .-` ".o o .O -.L N as. a -. a N en pt o .g "a. o Oa.a 9 ." ro! a o oa -L aNa o o oa p ."'a aNa! gg ¥ a e' :- g. o ¥¥ g vo sa or N ro N m va = pt N 5' Hg ¥¥¥. : ason g' = pt 6 A A # 6 e.-.ca gs B -. y.gg z m o m m G 69 E - ve. o { s g i g e ca o : ca 3g mff o st stm !¥ " :¥ 2¥ i' {- g tr. :t e. tr. "- tt. = . o ny 9 o g .? Sg =co o =co g ON gg. J co =o " gs s'mp pt e 6' M== v) tha 66 ,The Journal of School Education Research, 2007, Vol,19 Shkibetsuriver' '.7 Tekachiriver 2" '7e 26tht'5IE"e4a e 8sm ¥ ¥eeee cg4 e-e ' E tt 3 2 1 1 oo o O51015statio2nO Maruyamariver 7 Shinanoriver 7 g"6 no its5s e6 " m eNee t'5- eOe . 54o eooeee $4 e¥ Eo E3 3 2 2 1 1o oo o 510152e 2530Station $tation Yatariver 7 Muko river g g6 ts5Ere4 ts 5 g4 o E3 . 2 1 1 o o o 2 4 6 4station5 Station 8 o7 Kako river Iboriver sa'6 a26m :t5 ts5- :.4 g4E32 E3 ¥ ¥ 2 1 8 o o 2 4 6 8 10 o Station Station Kino river Shlrnantoriver 7 v7 g a26co g6 :t5 ts.5 -g4E32 .4 23 ¥ 2 1 8 o o 5 10 f5 Station Station Fig. 3. Distribution of mean teaf shapes of WittOW popuLation along river. For further exptanatio' n, see the {egend of Fig. 2. Predominance of' female and slender leaf in riparian willow populations 67 Shiribetsu river Tokaehiriver e'g 10 ¥ ege es Z-E8 ¥7 sE8 ts Åë7 ¥ .6 -o6d ¥ o=5 c5 = g4 eQ"ee E3g4 S3 2 2 "e 1 g o 5 10 o Statien 15 Station Shinanoriver Maruyamariver 10 10 va eg t9v o E8 E8tu if7o E7 e6ci vo6. - =5o =5 84c 84e E3 E3 2 se 2 1 to o o 10203040Station Station Yatariver Muko river 10 10 e 9 eg vE-e 8 es ÅëL-. 7 g7 "e' i 65 gs 6 es 432 eE g4 E3 2 eb-rtr'`2""-""-N.. 1 1 eo o 1 2 3 o 2 4 6 4station5 Station 8 Kakoriver iboriver to 10 m E 9 v= 9 oEs; e 87 E 8 s = 7 ts ts 65 o 6 g' o. c 5 c ovE 432 es 4 m E 32 1 1o o 4 6 4 6 8 Station Shimantorlver 10 Kino river 10 {9 g 9 es e 8 Åí7 Åë-s 7 ts 6 o 6 ' 2' s 6 = 5 i g4 o 4 E3 E 3 ¥ 2 2 ¥ 1 1 ¥ e o o 5 10 15 o 5 IO Station 1520Station Fig. 4. Distribution of mean numbers of fitaments of mate ftower atong river. For further exptanation, see the- tegend of Fig. 2. 68 The Joumal of School Education Research, 2007, Vol.19 Shiribetsuriver Tokaoh-- 1 8 1 mver ¥--o E O.9- E O.9 lj af O.8 O.8 co ca O.7 O.7 ¥ ¥ O.6 . O.6 "" O.5 O.5 . O.4 O.4 e O.3 ' O.3 O.2 02 O.1 O.1 o o o 1 2 3 4 5 6 7 sgiog{l,,ig2, 2 3 4 5 6 7 8 9 1011 12131415 ot Station Shinanoriver Maruyama.rlver ."o 1 e 1 e O.9 e O.9 lj O.8 s O.8 co O.7 "e O.7 O.6 ¥ . O.6 O.5 e -e O.5 . O.4 " O.4 e.3 . O.3 O.2 O.2 O.1 O.1 o o o 5 10 15 20 2 3 4 7 2530Station Ol 56Station Yata river Muko river ."o l ..e..1 E o.g e o.g X O.8 ts O.8 co O.7 ""'"'"'v------apt---. co O.7 O.6 O.6 ¥ O.5 O.5 O.4 O.4 O.3 O.3 O.2 02 O,1 O,1 o o o 1 2 3 statioA o 1 2 3 4 5 67 Station Kino river .9 1 Kako river ..o-1 :' O.9 SO.9 X o.g :O.8 co co O.7 O.7 O.6 O.6 O.5 O.5 O.4 . O.4 O.3 O,3 O.2 O.2 O,1 O.1 o o o 1 2 3 4 5 6 7 7 8 9 10 11 12 89Station Ol 234 56 Station ..o-1 Shimanto river g o.g 8x O.s O,7 O.6 O.5 ee O.4 O,3 O.2 O.1 o o 2 4 6 8 10 t2 14 16 18 20 Station Fig. 5. Distribution of sex ratios of witlow poputations atong river. For further exptanation, see the tegend of Fig. 2. Hokkaidodistrict Shin'etsudistrict 52,8A3 y=O.0254x+O,1606'r=+O.33 g.2.8t-.2.6A3 y=O.OO16x+1.t972 ."> -2.6m pt+O.08 ts2.4 bz4>V2.2 S-2.2 "-82 82 81,8 'g1.8 ee"Neeee .a1.6 .a1.6 1.4 1.4 1.2 ee eS 1.2 e ee g ¥ v e g O.4 O.4 oe O.2¥ O.2 s' g o o 8 4050 60 70809010g110120130140 4050 60 708090100110120130140 a tho Warmthindex Warmthindex E su o g pt Kinkidistrict NankiandShikokudistricts g 3 3 g 52.s>2.6.e g A y=O.0436x-3.5944 AEE2.s H .v" y=O.029x-2.5371 tr rt-+O.73 -2.6 po 6 g2.4 g2,4 r-+O.59 5' i-2.2 i-2.2 :¥ vR N' 'g1.832 -g1.882 5 ee e"ee q .a1.6 .a1.6 eee e' o 1.4 1.4 a 1.2 1.2 e"e 8v 1 e 1 g- . p O.8 O.8 5' See : O.6 O.6 eee"ee O.4 e O.4 oC e.2 02 ¥ o e o ' 90100 110120130140 405060708090100110120130140 4050607080 Warmthindex Warmthindex Fig. 6. Corretation between species diversity (H') of witLow poputations and Wl. The distribution area was dMded into 4 districts. The Hokkaido dis- trjct indudes the rivers of Shirjbetsu and Tokachj, and js inctuded jn the summer green vegetatjon zone. The Shjn-etsu djstrict indudes the Shinano river, and is tocated in the transient vegetation zone between the summer green and evergreen broad-teaved vegetation zone. The Kinki district inctudes the rivers of Maruyarna, Yata, Muko, Kakogawa and lbo. The Nanki and Shikoku districts inctudes the rivers of Kino and Shimanto. The tast 3 districts are inctuded in the evergreen broad-teaved vegetation zone. Linear egression tines and Pearson's corretation coefficients are atso shown. Da }g Hokkaidodistrict Shin"etsudistriot 765 /s" g2e8" y=O.Ol93x+3.8879 y=-O.0356x+72e58 .eae"eeet r-+O.51 !eu,s. pt-O.34 ¥ ee eeS . 4 4 3 3 2 2 g o el' tr 1 E su o40 o. ge 405060 7esogoloo 110120130140 5060 708090100HO120130140 g Warmthindex WarTnthindex 9 tu g 8 g¥ Kinkidistrict NankiandShikokudisbict = 7 pa ,k g2m6k gg g e O.O078x+2,B637 8 y= n y=O.0653x-2.6496e r:+O.31 -tr F-O.70 ¥8 2 t¥ .s .o 5 5 . < ) ee p 4 ¥ 4, ¥ eQ eee ¥ 6 3 3 ¥ e 2 21 1 o o 40 120130140 4050 120130140 so.6o7osogoloo11e Warmthindex 6o7osogeloo"o Warmthindex Fig. 7. Corretation between mean teaf shape of wiUow poputations and Wl. For further exptanation, see the tegend of Fig. 6. Hokkaidodisthct Sh;n-etsudistrict 103=o9EÅí8 .10 ey=-O.0684x+7.1316 Go9Ees y=O.O057x+1.3274 r=-O.18 -pt+O.28 "O- L,O" .7 e 7 2=6 dE6 Ere'5 se .. ."o5 4 4 v g 32ie o B eE 6' pt ,r g o o. oth e pt- o. 405060708090100MO120t30140 B Warmthindex Warmthindex pt m-. o" pt R o- Kinkidistrict ge NankiandShikokudistrict th .10 B' to9E y=O.029x-1.4183 y=O.0362x-2.5193 g pt+O.50 F+O.68 B, Åí8 eas8 g - ts e' v o7. o o97 o a =6sc E6 8v ru N s E05 p S5 5' : 4 - 4 32 e8 32 eeAe 1o 1o 40 5060708090100110120130140 405060708090100110120130140 Warmthindex Warmthindex Fig. 8. Corretation between mean number of fitaments of maLe ftower of wittow poputations and Wl. For further exptanation, see the tegend of Fig. 6. l Fg Hokhaidodistriot Shin-etsudistrict 1 'siO.9x o ol y=-O.O029x+O.7341 'go.g y=e.oeosx+o.so32 F-O.27 r-+O.12 x 8o.s oco O.8 O.7 O.7 O.6 . O.6 . .e.ee ee O.5 O.5 e O.4 O.4 . O.3 O.3 otrH O.2 O.2 tr O.1 O.1 fi pt- o o a 6e ge 4050 708090100110120130140 405060 708090100110120130140 g Warmthindex Warmthindex 9 ca g 8 g¥ Kinkidistrict NankiandShikokudistrict 5 ol'pEO.9xcOoo.B rv ol'peosxdiO.8 y=O.0 $ 28thx.is02868eee y=O,OO08x+O.5131 ee r==+O.13 .a. 8 .o O.7 O.7 . < O.6 eeeett.¥ O.6 g ¥eekee"e G O.5 O.5 O.4 e¥ O.4 . O.3 O.3 O.2 O.2 O,1 O.1 o e 405060708090100110120130140 40 5060708090100110120130140 Warmthindex Warmthindex Fig. 9. Corretation between sex ratio of wittow poputations and Wl. For further expEanation', see the tegend of Fig. 6. Predominance of female and slender leaf in riparian wirlow populations 73 ' DISCUSSION Hoklcaido district, but a positive significant correlation in the southern districts (Fig. 7). There also exists a negative ' Willows as pioneer plants in the early successional correlation between WI and the mean number of filaments stage in the Hokkaido district, but a positive significant correlation Willows have a wide range of distribution, from alpine in the other districts (Fig. 8). The transient stage can be grassiand in Hokl(aido to estuary of Shikoku and Kyushu. seen in the Shin-etsu district. If number of filaments of A total of 42 species are described in our country male flowers is one of indications of systematic relationship (Kimura 1989). Out of them, 12 species grow in alpine of willows, S. chaenomeloides and S. subfragilis seem to grasslands. The corresponding WIs are less than 15. A total be actual pioneers toward the warmer climates. The forrner of18 species inhabit in rather dry condition, including the species has several filaments from 3 to 5, and the latter alpine species. The remaining 24 species grow in riverside has 3. The remaining Salix species have 2 or 1. The former and lakeside, and are called riparian trees. All the willow species are seen only in mouth regions of rivers in the trees are deciduous, and the center of the distribution of southeastern area of our country (Fig. 1) (Ishikawa 1988). willows is expected to be in the summer green vegetation The jointed positive correlations are effectively indicative zone (Niiyama 1990). Actually, the maximum value of of an adaptive form of willow populations to the warmer H', 2.2, was observed in Tokachi river. The corresponding climatic niches. WIs are in the range from 45 up to 85. Some ripqrian willows A possibility of monoecious species with vegetative grow also in the evergreen broad-leaved vegetation zone. proliferation and the adaptation strategy of willows to These WIs are more than 85. 0ut of the rivers examined warmer frontier in the present study, all the STs of Shiribetsu river and A male of S. eriocarpa was never recognized in a total of Tokachi river are included in the summer green vegetation le39 individuals examined in this study, but the females zone. The upstream STs of Shinano river are also included were popularly observed only in the Kinki, Nanki and in the zone. The remaining STs are in the evergreen broad- Shikoku districts. The growing sites of this species are leaved vegetation zone. Therefore, Shimanto riVer seems to restricted to riverside but lakeside or even irrigation side. be in the transient stage between the major vegetation zones. Spreading of this species is impossible by usual fertilized Riparian condition is frequently suffered from disturbance seeds. A mixture of female and male flowers was tried to due to flood and freshet. Plants on riverside or floodplain examine in single individuals, but no evidence was gained. seem to be pioneer species in the early stage of succession However, the species was found to have a high ability of (Abe 1999).This is easily recognized even in the evergreen formation of adventitious roots from broken branches and broad-leaved vegetation zone, because of their deciduousness leaves. One of the highest value of O.64 was gained and life forms. The life forms of the willows in the zone (Table l). A single leaf or even a part of it works per- are MM in' Raunkier's criteria, and the disseminule form is fectly as an effective seed in this species. wnen this is the Di (Raunkiaer 1934; Numata et. al 1975). However, the case, the predominance of females has an advantage to actual life form seems to be like Th due to high vegetative that of males (Crawford et. al 1983; Takehara 1989;). growth induced regularly from broken branches and spread From a cross of European wiillow of S. vimin. alis, a wide leaves. These ecological traits are also the case in invaded range from extrerne female bias to extreme male bias took or naturalized plants (Shimizu 2003). They commonly have place in a homogeneous environment and in the absence a rapid growth after germination, plenty of seed production, of herbivores. Multiple gene system is possible to explain rapid proliferation by vegetative growth and so on. a sex determination in willows (Alstrdm-Rapaport et. al From the above point of view, the correlations 1997). Slender leaves more effectively fall into water among ecological characters appeared in Table 2 were re- streams, and reach to places to grow suitably. They are analyzed by dividing them into 4 local districts. There is also more easily broken into several pieces than round ones. no clear correlation between WI and H' in the Hoklcaido A close species of S. pierotti has the same tendency, and the and Shin-etsu districts, but there is significant positive cor- jointed sex ratio is O.84. This means that the frequency ,of relation in the Kinki and Nanki-Shikoku districts (Fig. 6). male plants is merely O.16. But the maximum of the The similar phenomena take place in the characters of leaf frequency of rooting from leaf was observed to be O.71 in shape and number of filaments of male flowers. There is S. pierotti. In addition, three mixed flower ears with female a fiegative correlation between WI and leaf shape in the and male organs were also found in a total of 1010 74 The Joumal of School Education Research, 2007, Vol.19 Tab(e 2.Corretation among ecotogicat conditions, systematic characters and species diversities. Ecoloicalcondition Sstematiccharacter Seciesdiversit Warmthindex Coldne$$index Riverbedgradient Meanno.filaments Meanleafshape No.ofspecjes Pielou'sinciex Shannon'sindex (WI) (Cl) (R.B.G.) (s) (J¥) (H') Sexratio +O.29"-t-O.23'rf).21" -.13-D.14 +o,eg-o.o4+o.o2 Warmthindex +O.95**-O.31"' **zz-O.14-O.25-O.25 Coldnessindex -O.22*-O.34"-O.34" Riverbedgradient Meanno.'filaments S-O.13 +O.42"+O.40"+O.46*' MeanIeafshape +O.39*'+O.47"+O.50" No.'ofspecies +D.62"+O.86*' Pielou'$index +O.90'" *Signifioant at 5% leveL "Significant at 1% level, ' individuals of the species, or O.O03 in frequency. 1-10 (in Japanese with English summary). The maximum frequency of the mixed flowers was ISHIKAWA S. (1983) Ecological studies of the floodplain observed to be O.Oll in S. chaenomeloides. The secondary vegetation in the Tohoku and Hokkaido districts, Japan. highest frequency was O.O04 in S. subfragilis. The sex ratios Ecological Review 20: 73-114. of these two speciesis are O.45 and O.41, respectively in this ISHIKAWA S. (1987) Ecological studies on the willow communities order. Therefore, the high productivities either by fertilized on the SatsUnai River fioodplain, Hokl(aido, with specieal reference seeds in single individuals or by vegetative growth seems to to the development of the Chosenia arbutifolia forest, Mem. be a compensatory mechanism to adaptation to the warmer Fac. Sci, Kochi Univ. Sen D (Biol.) 8: 57-67. niches for willow species such as S. chaenomeloides, S. ISHIKAWA S. (1988) Floodplain vegetation of the Shimanto subtfragilis, S. pierotti and S. eriocarpa. The jointed mecha- river in Shikoku, japan I. Arrangement of the main plant nism of above two factors seems to show the strategy of commimities developing on the bars in the lower course. Mem. adaptation of the willows to new frontiers. It really hap- 'Fac, Sci, Kochi.Univ. Sen D (BioL) 9: 25-31. pens in riparian willow populations at the Kinki, Nanki ISHIKAWA S. (1996) Characteristics of riverside Plants. In: and Shikoku districts of our country. Riparian environment and aguatic plants '(eds S, Okuda & Y. ' Sasaki) pp.116-139 (in Japanese). Softscience Co, Tokyou. ' REFERENCES KIMURA A. (1989) Salicacea. In: vaildflowers ofJapan, VVoody plants (eds Y. Satake. K. Hara, S. Watari & T, Tominari) pp. ABE S. (1999) Changes ' in species composition and life form 31-51 (in Japanese). Heibonsha, Tokyo. spectra of stands during the development of riparian forest in KIRA T. (1948) On the classification of vertical vegetation belts the Tanzaawa Mountains, central Japan. Japanese Journal of by warmth index. Agriculture in cold district 2(2):143-l73 (in Ecology 32: 247-257 (in Japanese with English summary), Japanese) . ALSTROM-RAPAPORT C,, LASCOUX M. & GULLBERG NIIYAMAU. K, (1987) Distribution of salicaceous species and soil (1997) Sex determination and sex ratio in the dioecious shrub texture of habitats along the Ishikari river. Japanese journa( of Salix viminalis L. Theor. Appl. Genet. 94: 493-497. Ecology 37: 163-174 (in Japanese with English surnmary). CRAWFORD R. M. M. & BALFOUR J. (1983) Female predomi- NIIYAMA K. (1989) Distribution of Chosenia arbutifolia and nant sex ratios and physiological differentiation' in arctic willows, soil texture of habitats along the Satsuriai river. Japanese journai Journal of Ecology 71: 149-160. of Ecology 39: 173-182 (in Japanese with English summary), GRANT M. C. & MITTON J. B. (1979) Elevational gradients in NIIYAMA K. (1990) The role ofseed dispersal and seedling traits adult sex ratios and sexual differentiation in vegetative growth in Åëolonizaion and coexistence of Salix species in a seasonally rates of Populus tremuloides Michx. Evolution 33(3): 914-918. fiooded habitat. Ecological Research 5: 317-331. ISHIKAWA S. (1980) Ecological studies of the floodplain willow NIIYAMA K. (1995) Life history traits of salicaceous species forests in the Hoklcaido district, Res. Rep. Kochi Univ. 29: 73- and riparian environment. Japanese Journal of Ecology 45: 78 (in Japanese). 301-306 (in Japanese). ISHIKAWA S. (1982) Ecologioal studies of the floodplain wil- NUMATA M., YOSHIZAWA N., ASANO S,, KUWAE[ARA Y,, low forests in the Tohoku district. Res. Rep, Kochi Univ. 31: OKUDA S. & IWASE T.(1975) PVeedflora ofJapan = Illustrated Predominance of female and slender leaf in riparian willow populations 75 by color. pp.406 (in Japanese), Zenkokunosonkyouikukyokai, Tokyo. PIELOU E. C. (1966) The measurement of diversity in different types of biological collections. Journal of 77ieoretical Biology 13: 131-l44. RAUNKIAER C. (1934) Life forms ofplants and statistical plant gTehOeiOcgy/argneanogn tphreessC,O(j/e.Cfitoerdd Pape"S of C¥Raunkiaer. pp. 64s, SAITO S. (1994) Classification and distribution of major willows in Hokkaido. Tree Breeding in Hokkaido 37(2): 24-33 (in Japanese). SHANNON, C. E. & WEAVER W. (1949) The mathematical theory of communicatio,n. pp. 117, Univrsity of Illinois Press, Urbana. SHIMIZU T. & CHIKATA F. (2003) On naturalized plants. In: Naturalized plants of Japan (ed. T. Shimizu) pp.11-39 (in Japanese). Heibonsha, Tokyo. TAK[EHARA A. (1989) Flowering size, fiowering age and sex ratio of willow populations along the Hirose River, nertheast Japan. Ecological Review 21: 265-275. (Reseived September 1, 2006; Accepted October 17, 2006)