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Home , Filipendula, Filipendula glaberrima, Filipendula multijuga, Filipendula palmata, Filipendula rubra, Filipendula ulmaria

植物研究雑誌 J. J. Jpn. Bo t. 69: 69: 29 0- 319 (1994)

Taxonomic Revision of the Genus Mill. ()

Ivan Ivan A. SCHANZER

Main Main Botanical Garden ，Russian Academy of Sciences Botanicheskaya 4 Botanicheskaya Str. ，Moscow ， 127276 RUSSIA

(Received (Received on J組 U 紅 Y 31 ， 1994)

百le genus Filipendula is taxonomically revised. 1t consists of 15 distributed in temperate areas of

血 eNorthern Hemisphere. Th e genus is divided into four sections defined on mo 叩hological and geographical

data.Filij フendula seems to have evolved mainly geographic through speciation during migrations and isolations

in in the late Miocene to Pleistocene. Th e main implied migration routes 台om North America to East Asia and farther farther westward via Himalaya and Siberia ，as well as distribution and variability pattems of the species ， and their relationships relationships are discussed. A key and a census of the Filipendula species are presen t.

The genus Filipendula Mill. consists of 15 species Materials and Methods of of perennial distributed over the m 吋or part of Th is revision is based mainly on the collections of the the temperate zone of the Northem Hemisphere. following Herbaria: E ，GH ，K ，LE ，MAK ，MHA ，MO ， Filipendula Filipendula was established as a separate genus from MOSP ，MW ，PE ，SHIN ，TI ，US ，VLA ，WTU. In total

Spiraea L. by Maximowicz (1879) ，who transferred it 1911 herbarium specimens hav~ been studied. from the Spiraeae to the Sanguisorbeae tribe ，印.ぷi e Th e field work was carried out in European Russia

Ro 凶so 凶id 白ea 鵠ei 泊nmodem 問sen 恥ce. Since then ， the position (1 98 6- 1991) and the Russian Far East (1 985 ，1987- of of the genus has been discussed repeatedl y (J uel1918 ， 1989). Seven species ，viz. F. ulmaria (L.) Maxim. ， F.

Sterling Sterling 1966 ，Baker and Baker 1967 ， Savile 1968) ， stepposa Juz. ， F. vulgaris 乱10ench ， F. angustiloba and now the majority of the authors places it in (Turcz.) Maxim. ， F. palmata (Pall.) Maxim. ， F. ， though in a separate tribe Ulmarieae. glaberrima Nakai andF. cα mtschatica (Pall.) Maxim. Taxonomic revisions of Filipendula have been have been studied in wild. Th e taxonomic transect carriedoutby carriedoutby Juzepczuk(1941 ，1955) ，Shimizu (196 1) method (Skvortsov 1968) has been practised to sam- and Sergievskaya (1 967) ， the latter work has re- ple populations of F. ulmaria ， F. stepposa ， F. mained unpublished. They suggested three different camtschatica ， F. glaberrima and F. palmata x F. treatments treatments of this genus ，each considerably discord- α ngustiloba hybrids. Th is method includes register- ant ant from each other in number of recognized species ing of several selected characters from each and pattem of infrageneric classification (Table 1). along an arbitrary profile through a local population This This paper represents an attempt to provide a new of a taxon under consideration to ascertain whether taxonomic treatment of Filipendula on the basis of there exist two or more discrete sympatric forms ， or a more comprehensive data and some new inte 中reta- continuous pattem of variability. tions tions of species distributional pattems. 1 have seen neither living nor herbarium of F.

-290- October October 1994 Joumal of Japanese Vo l. 69 No. 5 291

Table Table 1. A comparison of different classifications of Filipendula

Juzepczuk(1941 ，1955) Shimizu (1961) Sergievskaya (1967) Schanzer (here)

Isubgen.Hypogyna Isubgen.Hypogyna Isubgen.Hypogyna 1 sec t. Hypogyna 1. 1. F. occidentalis 1. F. occident αlis 1. F. occidentalis 1. 1. subgen. Aceraria 11 subgen. Ul maria 11 subgen. Aceraria sect. 1 sect. Schalameya 1 sec t. Schalameya 1 sec t. Aceraria 11 sec t. Schalameya 1. 1. F. kamtschatica 2. F. kamtschatica 2. F. kamtschatica 2. F. camtschatica 2. 2. F. glabra 3. F. glaberrima 3. F. koreana 3. F. glaberrima 3. 3. F. purpurea 4. F. yezoensis 4. F. auriculata 4. F. fomosa 4. 4. F. palmata 5. F. purpurea 5. F. glaberrima 5. F. multijug α 6. 6. F. multijuga 6. F. vestita 6. F. tsuguwoi 7.F.p α lmata 7. F. palmata 7. F. kiraishiensis 8. 8. F. rubr α 8. F. purpurea 11 11 sect. Albicoma 11 sect. Albicoma 11 sec t. Albicoma III sec t. Albicoma 5. 5. F. angustiloba 9. F. angustiloba 9. F. angustilob α 8. F. αngustiloba 6. 6. F. intermedia 10. F. intermedia 9. F. palmata 11. 11. F. rubra 10. F. rubra 11 11 subgen. Ul maria III sec t. Sessilia III sec t. Gymnocarpa 10. 10. F. vestita 12. F.formosa 11. 11. F. kiraishiensis 13. F. kiraishiensis 12. 12. F.formosa 14. F. tsuguwoi 13. 13. F. tsuguwoi 15. F. multijuga 11 11 subgen. Ul maria IV sect. Ul maria III subgen. Ul maria IV sec t. Filipendula 7. 7. F. ulmaria 14. F. ulmari α 16. F. ulmaria 11. F. vestita 8. 8. F. denudata 17. F. denudata 12. F. ulmaria 9. 9. F. stepposa 18. F. stepposa 13. F. stepposa 10. 10. F. megalocarp 。 19. F. megalocarp α 14. F. megalocarpa III III subgen. Eu イilipendula III subgen. Filipendula IV subgen. Filipendula 15. F. vulgaris 11. 11. F. hexapetala 15. F. hexapetala 20. F. hexapetala

tsuguwoi Ohw i. Filipendula tsuguwoi is an endemic 九1orphology of of Kyushu and Shikoku Islands in Japan. For mor- Underground organs Filipendula species are phology ，chromosomes and distribution 1 rely upon mainly shortly rhizomatous herbs with sympodially data data available from Ohwi (1954 ， 1965) and Shimizu branching rhizomes. This type of a life fo ロnlconsider (1961) (1961) publications. to be a primary one (Schanzer 1989a). Two groups of Scatter Scatter diagrams and dot maps have been used to deviate species ，however ，from this type (Fig. 1). analyse analyse all the herbarium material available. These has very peculiar and unusual methods proved to be very informative and provided among the rest of the species root tubers working as a the the main tool in studying pattems of geographical means of storage and vegetative propagation. variability variability and geographical distribution of characters North American F. rubra (Hill) Robins. ，and East in in most of the species. Asian F. palmata ，and F. angustiloba have long thin Chromosome counts were made in root tips of subterranean runners with annual accretion up to 10- seedlings seedlings of F. ulmaria ，F. vulgaris ，F. camtschatica ， 20 cm ，specialized for vegetative expansion and propa- F. glaberrima and F. palmata after Baker and Baker gation. Some of these species can rarely pos- (1967) (1967) with minor modifications. Voucher specimens sess rhizomes of the first unspecialized type. are are kept in MHA. Generati νe stalk All the species of Filipendula 292 292 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

have a ribbed hollow stalk ， with the only exception for F. vulg αris ，which has a solid one.

Le αif of Filipendula 訂 e interruptedly

p加natisected with palmately lobed terminal segments. 2 Leaves of different species ，however ，can vary sig- nificantly nificantly in size ， as well as in shape and number of segments (Fig. 2). The of F. vulgaris has a small (not larger than lateral lateral segments) 3-10bed terminal segment and nu- merous (usually 7-14 pairs) lateral ones. Other spe- cies cies usually have leaves with terminal segments larger

Fig. Fig. 1. Underground rhizomes of Filij フendula (schemati 目 than than lateral ones ，which are not so numerous. cally). cally). Scale bar = 1 cm. 1. Long subterranean runners (F. palmata ， F. angustiloba ， F. rubra). 2. Short rhizome Filipendul αrubra ， F. palm α ta andF. angustiloba ， typical typical for majority of the species. 3. Rhizome and root already already mentioned above for their peculiar rhizomes ， tubers tubers of F. vulgaris.

9

10 解

Fig. Fig. 2. Leaves and stipules of Filipendula. Scale bar = 1 cm. 1. F. camtschatica. 2. F. occidentalis. 3. F. F. glaber ァima. 4. F. multijuga. 5. F. ulmaria. 6. F. stepposa. 7. F. angustiloba. 8. F. palmata. 9. F. vulgaris. vulgaris. 10. F. kiraishiensis. October October 1994 Journa1 of Japanese Botany Vo l. 69 No. 5 293 have have leaves with terminal segments cleft into 7-11 parts. Ovaries contain 2 ovules ， one of which only deeply deeply cut lobes. Lateral segments (1-4 pairs) 訂 e develops into a seed. diameter varies among

(2)3-5 (2)3-5 lobed ，too. species from 6 to 15 mm. Flower features 訂 e usually The rest of the species have leaves with 3-5(7) rather constant and taxonomically significant mostly lobed lobed terminal segments and 1-6 pairs of lanceolate ， at a leve species l. Some of these characters 訂 e listed sometimes sometimes deeply se 町 ate ， but never lobed lateral below (Fig. 3). ones. ones. Differencies between these species in leaf shape Several more or less distinct types of shape are are not so easy to describe and in case of diagnostic can be distinguished within Filipendula. Long (ca. 5 importance importance they are mentioned in a key below. mm) narrowly triangular acuminate green of F. One more group stands out from the rest of the occidentalis have fairly seen midribs and often 1 to 2 species species for a peculiarity of its leaf venation pattern. 1t teeth. Presence of a sepal midrib is a unique trait of this consists consists of F. ulmaria ， F. stepposa ， F. meg α locarpa species ， and 1 suggest it to be plesiomo 叩hic ， because Juz. ， and F. vulg αris ，whose terminalleaf segments of reduction of a midrib in all other species ，which can have have first pairs of lateral veins always being SUf- possess this character only as a haphazard variation. rounded rounded by lamina tissue but never stripped at the Green sepals of similar size and shape ， but usually base. base. On the contrary ，first pairs of terminal leaf without midribs and teeth ， are typical for F. vestita ， F. segment segment lateral veins of all the other species are stepposa ， and F. megalocarp α. These taxa ，however ， usually usually stripped at the segment base. can possess also more short triangular ， tapered or Stipules Stipules are more or less auriculate and rounded or rounded sepals not always green in colou r. Sepals of elongated elongated in outline. Stipules of F. occidentali ・'s (S. the rest of the species are usually yellowish or brown- Wats.) Wats.) Howell ， F. kiraishiensis Hayata ， F. formosa ish ， sometimes pu 叩le. Sepal shape is usually of no Nakai Nakai andF. multijuga Maxim. lack usually auricles. taxonomic significance due to its variability even 1n 1n the last two species they are lanceolate-oblong in within the same . outline ，obtuse ， and often membranaceous. This strong Filipendula are orbicular to ovate in outline ， character character enables determination of these species even with smooth to slightly unevenly toothed margins ， by one stipule. The stipules of F. glaberrima and F. X and usually with distinct na 町 ow claws. They are purpur ωMaxim. are also remarkable ， being greatly coloured white to slightly yellowish ， often with pink reduced ， narrowly triangular ，usually drying up and to pu 叩le spots when being in buds ，rarely pink or fragmenting fragmenting after flowering. pu 叩le (e.g. ，in F. multijuga ， F. rubra ， F. Xpurpurea ，

Inflorescence Inflorescence The genus Filipendula is character- F. X auriculata (Ohwi) Kitam. et Murata ，and some- ized ized by its unique inflorescence - an anthela -a times F. gl αberrima). corymbiform corymbiform panicle with a shortened central axis Two species with unusual features are worth and and lower branches several times surpassing upper special mentioning. Petals of F. multijug αlackclaws. ones ones (Fig. 3). Bracts and bracteoles are always lack- Each petallinks to the margin by a very mg. n訂 row base which seems to be a reduced claw. Such Flower Fl owers of Filipendula have a saucer- claws reduced occur haphazardly in some other spe- shaped shaped (rarely nearly flat) hypanthium with 5 sepals ， cies as a variable character state ，e.g. in F. ulmaria. 1n 5 petals ， 15-25 ， and an apocarpous gynaeceum F. multijuga this character is fixed and seems to be with with 2 to 15 free carpels. The only exception is F. apomo 叩hic. vulgaris ，which perianth consists of 5 to 7， mostly 6， Filipendula occidentalis has large petals without 294 294 植物研究雑誌第69 巻第5 号 平成 6 年 10 月 claws ，linking to the hypanthium by broad base with convex towards achenes maturing because of sinking many veins getting into a petal instead of one in the of its margins down. The part central of a hypanthium ， rest rest of the species. Such character is unique within the a torus ， enlarges forming a small ca 中 ophore. This genus and seems to be plesiomorphic. tendency is characteristic of F. vulgaris ， F. stepposa ， A hypanthium in Filipendula is usually more or F. ulmari 孔F. megalocarpa ，and F. vestita ， though in less less saucer-shaped ，though its depth can vary consid- F. megaloc α rpa and F. vestita a ca 中ophore can be erably erably from nearly f1 at in F. occident αlis to rather absen t. Sometimes it can be weakly developed also in deeply deeply concave in F. vulgaris and F. kiraishiensis. F. angustiloba. This character seems to be apomorphic.

The last character state 1 believe to be apomo 中hic in A number of carpels varies usually from 5 to 11. F. accordance accordance to Shimizu (1961) and Kalkman (1988) ， vulg αris and F. stepposa have up to 15 carpels and ， who stated an increase in hypanthium concavity to be vice versa ， F. multijuga has only 4， rarely 5. South a general trend in Rosaceae. Korean F. formosa possesses 1-3 carpelsonly.

In In several species a hypanthium becomes f1 at to An apoc 訂 pous in fruit Filipendula -consists of

iQ Vハ 7

16 16 4 3 2

Fig. Fig. 3. Generative parts of Filipendula. 1. Inflorescence of F. palmata. Scale bar =1 cm. 2. 2. F. ulmaria peta l. Scale bar =1 mm. 3. F. multijuga peta l. 4. F. occident αlis peta l. 5. Hypanthium Hypanthium and achene attachment in F. cα mtsch αtica.Scalebar= 1 mm.6.Hypanthium and and achene attachment in F. vulgaris. 7-16. Achene shape and pubescence. Scale bar = 1 mm. 7. F. camtschatica. 8. F. glaberrima. 9. F. kiraishiensis. 10. F. palmata. 11. F. multijuga. multijuga. 12. F. angustiloha. 13. F. vestita. 14. F. stepposa. 15. F. ulmaria. 16. F. vul- gans. gans. October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 295 several several achenes. An achene usually has a stipe which covered with such pubescence. can be more or less reduced and placed at the base of Pubescence is usually rather variable within a an achene or shifted to some extent towards its ventral species except on achenes. The latter covering is quite suture. suture. An achene can also be more or less twisted. stable and thus deserves special mention due to its These characters are usually rather peculiar for each diagnostic significance. There are 4 types (Fig. 3): of of the species and allow to distinguish most of them 1. Ciliate hairs along achene sutures occur in F. (Fig.3). (Fig.3). occidentalis ， F. camtschatica ， F. glaberrima ， F. Pubescence Pubescence Above-ground offshoots of Filipendula pα lmata and F. vestita. The first two species also are are usually more or less hairy with simple unicellular ， possess sparse appressed hairs on the lateral sides of variously variously shaped trichomes. Five most usual types of the achene. the the latter are listed below ，though these differencies 2. Appressed hairs scattered throghout the achene are are not absolute and various transitional states can surface ，more densely along sutures ， are characteris- occu r. tic of F. vulgaris. 1. 1. Short (0.05 to 0.2 mm) squarrose hairs. They are 3. Sparse ciliate or sinuate pubescence occurs in very very stiff and dense on stems ，petioles ，inflorescence the upper part of the achene ventral suture in F. branches ，pedicels ，and sometimes leaf veins of F. stepposa and F. meg αlocarpa. This character appears camtsch αtica ，F. formosa ，and F. occidentalis ， often also in some of F. ulmaria and F. rubra appressed appressed forward in the last species. F. palmata and which normally have glabrous achenes. F. F. rubra bear similar ， but thinner and sparse hairs. 4. Glabrous achenes are found in all the other 2. 2. Long (0.5 to 2.0 mm) squarrose tortuous hairs. species. Among them only F. multijuga sometimes They are stiff and dense on stems of F. camtschatica possesses ciliate achenes in the northem part of its and sparse in F. rubra. More thin ones often densely range. These plants are known as var. ciliata. cover cover leaf undersurface veins in F. camtschatica ， F. glaberrima ， F. rubra ，and rarely F. ulmaria. Chromosome numbers 3. 3. Long (0.5 to 1.5 mm) sinuate hairs. They form The basic chromosome number in Filipendula is appressed dense to sparse pubescence on leaf x=7 (Baker and Baker 1967). Reports ofx=8 are now undersurface undersurface veins in F. rubra ， F. occidentalis ， F. considered as artifacts or mistakes (Baker and Baker camtschatica ， F. glaberrima ， F. kiraishiensis ， F. 1967 ，Love and Kjellqvist 1974). The following data formosa ， F. multijug α，and F. angustiloba. on chromosome numbers in Fil 伊endula are collected 4. 4. Long (0.5 to 1.5 mm) sinuate to curly thin hairs from literature and my own counts. forming appressed cobwebby pubescence on stems of Filipendula ulm α ria and F. vulgaris are diploids F. F. vestita ， F. megalocarpa and F. stepposa. Similar throghout their ranges with 2n=14 (Wulff 1938 ，

but but more squa 町 ose and sparse pubescence covers Turesson 1938 ，Polya 1949 ，L りve and L 凸ve 1956 ， stems stems and inflorescence branches of F. kiraishiensis Sorsa 1962 ， Gadella and Kliphuis 1963 ，Baker and and probably F. tsuguwoi. Baker 1967 ，Laane 1969 ，Kalda and Roos 1973 ，L りve 5. 5. Curly hairs forming sparse to dense whitish and Kjellqvist 1974 ，Kartashova et al. 1974 ， tomentose tomentose pubescence on leaves undersurface of F. Lempiainen 1978 ， Krogulevich 1978). These data vestita ， F. stepposa ， F. meg αlocarpa ， F. ulmaria ， F. cover the ranges of both species from the Atlantic palmat αand， F. angustiloba. Pedicels and inflores- Ocean to Altai ， East Sayan mountains and West cence cence branches of the first three species are also Siberia. My own data from central Russia (neighbour- 296 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

hoods hoods of Moscow and Tarussa) coincide with them: tetra- and hexaploid cytotypes (2n=28 ，42 -4 4: 2n=14 in both species. Sokolovskaya 1966). No further species have been A more complicated situation exists in F. investigated to date. camtschatica. camtschatica. The literature data are contradictory: 1t would appear difficult to draw any conclusion Funabiki Funabiki (1 958) revealed 2n=14-18 in plants of from such incomplete data. It has been. seen ， that Hokkaido ，J apan. Later Sokolovskaya (1960) counted diploid species: F. ulmaria ， F. vulgaris ， F. multijug α 2n=28 in plants of Sakhalin and 2n=28 ，42 -4 4 in and F. tsuguwoi ， possess a wide range of evidently plants plants of Kamchatka (Sokolovskaya 1963 ， apomo 中hic characters such as dioecoey ， tomentose Sokolovskaya Sokolovskaya et al. 1985). Gurzenkov reached the pubescence of leaves ， glabrous fruits ，etc. ， while same results (Stepanova et al. 1973). He stated that species with several polyploid cytotypes lack usually tetraploids tetraploids (2n=28) have hispid stems and leaves ， such characters. 1 believe that it is probable that whereas whereas hexaploids (2n=42 -4 4) have glabrous ones. morphological evolution in Filipendula has occurred This This has been accepted by a number of authors mainly at the diploid level ， while the significance of (Sokolovskayaeta l. 1985 ，Morozov andBelaya 1988). polyploidy is restricted. M Y own counts revealed a diploid number (2n= 14) in plants plants ofKunashir and Shikotan Is1ands ， and southem Distribution Sakhalin Sakhalin (S.-Sakhalinsk and Dachno 邑); a tetraploid Filipendula is distributed in the temperate zone of number (2n=28) was found in plants of central Sakhalin Eurasia and North America and also in the Himalaya (Roshchino (Roshchino and Tymovsk). 1n Nevelsk (southem and Yunnan Province of China. The species form Sakhalin) Sakhalin) both cytotypes occurred. It appears that three more or less distinct distributional groups shown three three cytotypes ，viz. di- ，tetra- ， and hexaploid ，exist in Figure 4. within within F. camtschatica. Diploids are distributed in the The first ，“ Pacific" ， group includes species with southem part of its range ，tetraploids in the central and ranges situated in the Pacific monsoon climatic zone. the the northem ， and hexaploids in the northem part only. The ranges of F. camtschatica ， F. glaberrima and F.

A thorough mo 叩hological study has found no co 汀 e- multijuga stretch along the Asian coast and the co 町 e- lation lation between the hairiness types and ploidy levels. sponding island arc ， whereas F. formosa ， F. tsuguwoi All All the types of pubescence occur throughout the F. and F. kiraishiensis are local endemics with ex- camtschatica camtschatica range. tremely restricted ranges in the same area. F. Filipendula Filipendula glaberrim α，a related species from occidentalis is an endemic of the Pacific coast of Primorsky Primorsky Province (Primorsky of Krai) Russia and N orth America. Korea ， apparently has two cytotypes: 2n= 16 Species of the second ，“ Continental" ， group are (Sokolovskaya (Sokolovskaya 1966; “ F. kore α na" from the south of distributed in the continental climatic areas of Eura- Primorsky Primorsky Prov.) and 2n=28 (Gurzenkov 1973: “ F. sia. Their ranges occur in a latitudinal direction from koreana" koreana" from the south of Primorsky Prov. ，Lazo the Atlantic coast of Europe to East Siberia and the pass). pass). My own counts from the same region (Khassan Yunnan mountains. These species are F. ulm αria ， F. Distr. ， Ryazanovka) show 2n=14. stepposa ， F. megalocarpa ， F. vestita and F. vulgaris. Two endemic Japanese species ， F. multijuga and The remaining three species form their own distri- the the dioecious F. tsuguwoi ， are diploids with 2n=14 butional group ，which is intermediate to some extent (Shimizu (Shimizu 1961). between two above. Two Asian species ， F. palmata Filipendula Filipendula palmat αfrom Primorsky Prov. has and F. αngustiloba ， are distributed in the Far East October October 1994 Journal of Japanese Botany Vo l. 69 No. 5 297

Fig. Fig. 4. Distribution groups of Filipendula. 1. ‘Continental' group (sec t. Filipendula). 2. ‘Subcontinental' group (sect. (sect. Albicoma). 3. ‘Pacific' group (sec t. Schalameya and sec t. Hypogyna).

penetrating penetrating deeply int o. the c o. ntinent as far as East pict o. grams (Fig. 5) sh o. ws ac o. incidence o. f the de- Siberia Siberia and Mongolia. Fil 伊endula rubra inhabits the fined sections with the ab o. ve mentioned distribu- Atlantic Atlantic regi o. ns o. f N o. rth America as far as the Mis- ti o. nal groups (see Fig. 4). Such co. incidence makes it sissipp i. Th us they dem o. nstrate an example o. f the p o. ssible to. reg 訂 d them as natural groups. So. me classical classical di 司uncti o. n between East Asia and Atlantic speculati o. ns ab o. ut their p o. ssible evoluti o. n are pre- N o. rth America. sented bel o. w.

Filipendula Filipendula occidentalis' is a na 町o. W endemic o. n Analysis Analysis of taxonomic and the Pacific co. ast o. f N o. rth America. It is no. do. ubt a phylogenetic phylogenetic relationships relic species ，alm o. st co. mpletely plesi o. m o. rphic in its Tax o. no. mic and phyl o. genetic assumpti o. ns were m o.叩 ho. lo. gical characters. Shimizu (1961) treated it made o. n the grounds o. fc o. mparative m o.中 ho. lo. gyand as a separate subgenus. 1 treat F. occidentalis as a ge o. graphy ，especially o. n the evidence o. f the ge o.- m o. n o. typic secti o. n cl o. sely related to. sec t. Schalameya graphic graphic distribution of characters. Th e results of the and very possibly ancestral to. it ， or to. the whole genus. m o. rph o. geographic analysis are visualized with the Secti o. n Schalameya is rather c o. mplicated. As seen help help o. f pictograms. Eight characters being ch o. sen to. in Table 3， the species o. f this secti o. n are related to. divide divide the genus int o. sections are sh o. wn in Table 2. each o. ther in a chain -like manner. There are tw o. m o. re The resulting pict o. grams are sh o. wn in Table 3. Fo. ur cl o. sely related groups o. f species: F. camtschatica and maj o. r subsets 訂 e seen. Secti o. ns Albic o. ma and F. glaberrim αin the no. rthem part o. f the secti o. n range Filipendula Filipendula are defined rather distinctly ， while the (Fig. 6) ，and F. tsuguwoi and F. kiraishiensis in the species species in sec t. Schalameya are related in a chain-like so. uthem part o. fi t. They are linked by tw o. species ， F. manner. manner. Th e sect. Hyp o. gyna is m o. no. typic ， and m o. st formosa from So. uth K o. rea ，and F. multijuga from similar similar to. sec t. Schalameya. Mapping o. f these so. uthem Japan ，which seem to. be cl o. sely related to. 298 298 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

Table Table 2. Characters and character states used in the Table 3 pictogram for sectional subdivision of Filil フendula

well well poorly Character Character deve- deve- absent loped loped loped Leaves Leaves tomentose beneath 。一 。 2 Underground runners 〈プ 。 3 Lateralleaf segments dissected 占 。 to to 2-5 lobes

4 Sepals inside appr 回l sedly hairy 、コ む 。 5 Achenes ciliate ， 一-0 。 。 (+)ー throughout hairy ←-0 (+) 6 Development of achene stipe 。 ρ メコ 7 Carpophore development ? ? 。 8 Achenes twisting 。 • ct

each each other. Together they form a range of vicarious species and hybrids are undoubtedly close to each

species species evidently differentiated in this territory dur- other. It seems co 町 ect to unite them into one section ， ing ing numerous migrations and isolations on islands distinguishing more closely related groups as subsec- and high mountains in late Tertiary and Quarternary tI ons or senes. (Golubeva 1982). It is quite possible that secondary Section Albicoma is well defined ， though F. rubra hybridization hybridization also could take place in the past as it seems to be transitional to some extent between sec- does does now in central Japan where ranges of F. tions Albicoma and Schalameya. It resembles F. camtschatica camtschatica and F. multijuga overlap (Fig. 7). The multijuga in petal shape and pink colour ，and F. resulted resulted hybrid F. x auriculata possesses an unstable camtschatica in sepal interior appressed pubescence combination combination of diagnostic characters and has been a and hispid stem pubescence. The peculiar leaf shape ， rather rather confusing taxon for a long time. It was sup- long subterranean runners ，and the form of the achene posed to be a separate species and a possible ancestor bring F. rubra close to Asian F. palmata and F. to to F. pwpurea known from garden culture only in angustiloba. Such a transitional position for this spe- Japan Japan (Maximowicz 1879 ，Ohwi 1965 ，Shimizu ， cies points out to its relative antiquity and is in good 1961). 1961). The analysis of its distribution and characters concordance with Kalkman' s of hypothesis (1988) an leaves leaves no doubt that it is a hybrid between F. American origin for the genus. The di りunction be- camtschatica camtschatica and F. multijuga. As to F. purpurea tween F. rubra and its Asian relatives F. palmata and itself ，1 believe it to be a horticultural hybrid estab- F. αngustiloba (Fig. 8) can be dated to the Middle lished lished by J apanese florists a long time ago. Filipendula Miocene ，when the Bering Bridge existed in temper- cα mtschatica ， F. multijuga ，and F. glaberrima could ate climatic conditions before sinking at the end of be its possible parents. All the above mentioned Miocene (Petrov 1976). October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 299

Table Table 3. A pictogram of Filipendula sectional subdivision

Hybrid Hybrid Pictogram Pictogram Section Hybrids Species Species plctogram O .角同」 F.occidentalis F.occidentalis 一¥。一、一七F出 2 F. camtschatica

3 F. gl α，'berrima 〈 とり F. x auriculata LF 同冨〈叫〈出 4 F.formosa こむ F. xpurpure α

5 F. multijuga -b um 6 F. tsuguwoi -)J や一ザ一#一 7 F. kir αishiensis

〈 8 F. rubr α， -40UHmHJJ一任 9 F. angustilob α ι F. x intermedia 刊

10 10 F.palmat α， トケ一ナ一全ナ一ヨ 11 11 F. vestita 〈凶己凸 12 12 F. megalocarp α Z 13 13 F. ulmaria 同缶、同比

14 14 F. stepposa

15 15 F. vulg αris

Fig. Fig. 5. Distribution of characters of the sections over the genus range. 1. Sec t. Schalameya. 2. Sec t. Albicoma. 3. 3. Sec t. Filipendula. 4. Sec t. Hypogyna. 300 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

and and some of its tributary valley meadows. andF.α ngustiloba are undoubtedly the clos- est est relatives ， and it is striking that in all previous treatments treatments of Filipendula (see Table 1) they were

sep 訂 ated to different sections on the formal grounds of of differences in achene pubescence and stipe length: F. F. palmata has ciliate stipitate achenes ，and F. angustiloba angustiloba has nearly sessile glabrous ones. Sec t. Filipendula is also well defined within the genus. genus. Similar to sec t. Albicoma ，where the di りunc- tion tion between American and Asian representatives exists ，it contains a species with a disjunct relic range ， F. F. vestita. Th e latter resembles some members of the sec t. Schalameya and Albicoma in having ciliate shortly shortly stipitate achenes ，stripped basal leaf veins ， and ， sometimes occurring in Yunnan populations ， plants plants with long subterranean runners. In other char- acters acters it is more close to sect. Filipendula. The range of of F. vestita (Fig. 9) points to a migration route of this species species from China to the West via the “Himalayan corridoor" corridoor" (Tabata 1988). 百lI s migration probably happened at the end of Miocene to the middle of Pliocene ，when the “ Himalayan_ corridoor" existed. Th e F. ulmaria sensu lato group (including F. megalocarpa ， F. stepposa andF. ulmaria sensu stricto) seems seems to be derived directly from F. vestita in the end of of its westward migration. Th e implied phylogeny of F. F. vestita - F. ulmaria sensu lato group has been viewed in detail in a separate paper (Schanzer 1989b). Fig. Fig. 6. Distribution of the sec t. Schalameya species. 1. F. This group represents a series of closely related vi- camtschatica. camtschatica. 2. F. glaberrima. 3. F. multijuga. 4. F. carious carious species (Fig. 9; Table 2). Filipendula formosa. formosa. 5. F. tsuguwoi. 6. F. kiraishiensis. megalocarpa is the closest to F. vest If， α， possibly derived derived from the latter during the migration from the The Asian members of sec t. Albicoma ， F. palm α ta Himalaya to Asia Minor ，where it st il1 persists ， via the and F. angustiloba ， share a number of common ch 紅- Kopetdagh and Elburz Ranges ，where it is now ab- acters ， are distributed in vicarious ranges (Fig. 8) ， and sen t. Similar stem cobwebby pubescence ， usually hybridize hybridize in the overlap area. Their hybrids ，known as more or less developed c訂 pophores ， and achenes F. F. intermedia ， are typically transitional between the with sinuate to ciliate pubescent sutures ventral bring p訂 ental species. They occur in the wild in morpho- them closely together. Filipendula stepposa and F. logically logically heterogenous hybrid swarms in the Amur ulmaria sensu stricto could be derived from F. October October 1994 Journal of Japanese Botany Vo l. 69 No. 5 301

Fig.7. Fig.7. Character distribution of F. cα mtschatica ， F. multijuga and their natural hybrid F. x uriculata αuriculata in Jap 加.

Characters Characters Character states

Petal Petal colour white 5 pink 。 2 Petal shape even 。 uneven uneven や ~i~::~;~s ? 3 Achenes number 3-4 。 4- 5 べコ 6-8 -0 4 Achenes pubescence glabrous 。 ciliate • 5 Sepal interior glabrous 。weak def 帥、 pubescence pubescence モコ

6 Stipule shape auriculate ，。 ashuorid culate ，u( ¥ lacks long long auricle c¥ 7 Petiole hirsute 。 unevenly glabrous 0- pubescence pubescence hirsute 。一 302 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

Fig. Fig. 8. Distribution of the sec t. Albicoma and Hypogyna species. 1. F. occidentalis. 2. 2. F. rubra. 3. F. palmata. 4. F. angustiloba.

megalocarpa megalocarpa during the northward migration of the somewhat dwarf rosette habit and sharply and

latter latter from Kopetdagh to the Urals ， where the first deeply se 町 ate leaves. Filipendula ulm αrz 仏 on the findings findings of F. ulmaria fossil achenes were made in contrary ， has the same habit as F. megalocarp α， but Upper Pliocene strata (Dorofeev 1965). Later ，at the lacks achene pubescence and other characters shared Pleistocene Pleistocene and Holocene ， F. ulmaria achenes be- with F. vestit α. The implied relationships and routes come a usual component of foss i1 assemblages in of migrations of these species are schematically pic- European European Russia and West Siberia (Dorofeev 1965 ， tured in Fig. 10. 1977 ，Katz ， Katz and Kipiani 1965 ， Velichkevich Th e position of F. vulgaris is a separate question.

1982 ，Koshkarova 1986). This co 汀 esponds with the All the previous monographers regarded it as a sepa- implied implied expansion of F. ulmaria and F. stepposa to rate subgenus. lndeed ，F. vulgaris possesses anumber Europe and Siberia during the Pleistocene and of characters quite unusual within Filipendula ，e.g. ， Holocene. Filipendula stepposa resembles F. root tubers ，multifid leaves ，solid stems ，5 to 7， mostly meg αlocarp αin having sinuate pubescence along 6 ，petals and sepals instead of 5 in all species other ， ventral sutures sutures ventral of the achenes ， and cobwebby pubes- and achenes that are hairy throughout their surface. At cent cent stems ， but differs strikingly from the latter in first glance this species seems to have no close rela- October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 303

Fig. Fig. 9. Distribution of the sec t. Filipendula species. 1. F. vestita. 2. F. megalocarpa. 3. F. stepposa. 4. F. ulmaria. ulmaria. 5. F. vulgaris.

Wacm 柑 d ):500 開。∞ IBlcM 5OO~.M) 5O{ 1 0 島国 間関 山'" 2創珂附

Fig. Fig. 10. The implied Tertiary-Quartemary migration routes of the ser. Ulmariae species. Modem distribu- tion tion ranges and achene outline are given. 1. F. ulmaria. 2. F. stepposa. 3. F. megalocarpa. 4. F. vestita. Land outlines shown in bold line correspond with lower to middle Miocene (After V. M. Sinitsyn ，1965 ， Drevnie Drevnie klimaty Evrazii (Ancient climates of Eurasia) ， part 1，Nauka ，Leningrad). 304 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

tives tives arnong other Filipendula. St i1l there exist several derived. So he supposed that their predecessor should

traits traits indicating to possible relationships of F. vulg αr 臼. be distributed in the Arctic region in the Tertia 守， The carpophore developing after flowering in F. from where it migrated to the south. vulgaris vulgaris is even larger than in any other species. This All the most primitive or plesiomorphic species of very very strong character bringsF. vulgaris close to theF. t. Sec Hypogyna and Schalameya are now distributed

vestita vestita - F. ulmaria sensu lato group. The achene in 訂 eas ofhumid ， and especially monsoon ，climate. It

shape shape is generally the same as in the latter group seems possible th 剖 the origin of Filipendula was

except except of twisting. Moreover ， achenes are almost connected with 訂 eas of such climate on the Pacific sessile sessile and attached to the ca 叩ophore at their ventral coast of America and/or Asia. t. Sec Albicoma and sutures ，by what it looks like a continuation of the Filipendula originated while spreading tocontinental

stipe stipe reduction trend in the above group. The ranges 訂 eas of Eurasia and America. The sec t. Filipendula of of the latter and F. vulgaris almost coincide ， and this migration route lay from China via the Himalaya to is is another argument in favour of their affinity. As for Asia Minor and the Urals ，from where it spread above above mentioned unique of characters F. vulgaris ， westward to the Atlantic Ocean ， and eastward to the

some of them ， such as root tubers and 柑 ongly dis- Lena River and Lake Baika l. Similarly the Asian

sected sected leaves ，紅e evidently adaptive to the dry habi- members of sect. Albicoma migrated to the west via tats tats of this species. The achene hairiness is probably north China and Siberia and reached the Enisey River. derived derived from a ciliate one: appressed flat are hairs is either the species ancestral to much more dense along sutures than on lateral sides of sec t. Albicoma and Schalameya ， or the result of a an achene. Similar hairiness is common in F. separate eastward migration from the hypothetical camtschatica camtschatica and ，especially ， in F. occidentalis. ancestral area at the Pacific coast of North America.

Unstability Unstability of the number of perianth p紅白 is no less remarkable remarkable and seems to be certainly a derived state. A key to the species of Filipendula Generally Generally speaking ， F. vulgaris seems to be a strong la Root tubers present; leaves with numerous lat- deviation deviation from the main pass of the F. vestita - F. eral segments (7-12 pairs); perianth of 6 (rarely ulmaria ulmaria sensu lato group evolution ， connected with 5 or 7) parts; achenes appressedly hairy through- the the adaptation of certain ancestral populations to out out the surface …-… H ・H ・-…...・ H ・...・ H ・..F. vulgaris much more severe dry habitats ，maybe via Quantum b1 Root tubers absent; leaves with few lateral seg- evolution evolution (Grant 1981). ments (1 -6 pairs) or them; without perianth of 5

According According to Kalkman (1988) ，Filipendula has p訂 ts; achenes hairy not throughout the surface or originated originated from South America ，where no representa- glabrous tives tives orrelatives ofthe genus exist flOW ， then migrated 2a Rootstock with long subterranean runners; lat- to to North America and Eurasia via the Bering Bridge. eral segments of middle cauline leaves deeply Such a conclusion has been reached by the analyses of palmately cleft into 3-510bes ，terminal segment distribution distribution of several non-related Rosaceae genera. cleft into 7-11 lobes Shimizu Shimizu (1961) considered F. occidentalis and N orth 3a Carpels and achenes ci1i ate; upper part of stem Asian Asian species of the section Schalarneya sensu Shimizu and inflorescence branches glabrous ， or shortly

(1961) (1961) to be the most primitive ， and south- and squa 町 ose pubescent (hairs ca. 0.05 mm) westward westward distributed species of the sections Albicoma ， 4a. Leaf segment lobes are not narrow linear ， Sessilia Sessilia and Ulmaria ， as well as F. vulgaris ， to be always greytomentose beneath ，at least at up- October October 1994 Jouma1 of Japanese Botany Vo l. 69 No. 5 305

per per leaves ...・ H ・....・ H ・H ・H ・-… H ・H ・-… ..F. palmata als pink ， with reduced claws .…

4b Leaf segment lobes 訂 en 訂 row linear and/or … F. multijuga v紅 . ciliata lack lack tomentose pubescence beneath .… 8c Sepals shortly appressedly hairy inside ・…… F. x intermedia 9a Inflorescence branches and pedicels

3b Carpels and achenes glabrous densely ，shortly ，bristly ，squa 町 osely pu-

5a 5a Leaf segment lobes na 町 ow linear ， with or bescent; petals white … ..F. camtschatica

without without tomentose pubescence beneath; in “ 9b Inflorescence branches and pedicels gla- florescence florescence branches sparse tomentose with brous; petals pink or pu 叩le ， at least in curly curly hairs; petals white ， with even margins; buds

sepals inside inside sepals glabrous ...・ H ・....F. angustiloba 10a Petals in buds pink ， then white ， with even

5b Leaf segment lobes are not n訂 row linear and margins; leaves hairy along veins be-

lack lack tomentose pubescence beneath; inflores- neath with squa 町 ose long hairs to almost cence cence branches glabrous; petals pink ， with glabrous; stipules small ， not auriculate ，

unevenly unevenly toothed m 訂 gins; sepals appressedly narrowly triangular ， often drying up and

pubescent pubescent inside '"・ H ・.....・ H ・..…...・ H ・.F. rubr α fragmenting afterflowering; achene stipe 2b Rootstock short ， without runners; lateral seg- shorter than one third of its fulllength .. ments ments of middle cauline leaves lanceolate ，some- .. F. glaberrima

times times deeply se 町 ate ， but not cleft into lobes ， 10b Petals pink to purple ， with unevenly rarely rarely absent; terminal segmentcleftinto 3-5(7) toothed margins; leaves hairy beneath

lobes lobes along veins with sp 町田 appressed hairs; 6a 6a Sepals with midribs; petals white ，clawless ， achene stipe equal to or longer than one with with broad base; petioles and inflorescence third of its fulllength branches branches appressedly bristly pubescent; lla Petals pink; stipules semiorbicular ，

achenes achenes densely ciliate ...・ H ・.F.occidentalis auriculate; leaf lateral segments in 1-2

6b Sepals without midribs; petals white ， pink or pairs '"・ H ・.....・ H ・.....・ H ・..… F. x auriculata purple ， with claws (if claws are secondarily 11 b Petals pu 中le; stipules small ， without reduced ， the petal base remains narrow); auricle ， narrowly triangular ， often dry- petioles petioles and inflorescence branches variously ing up and fragmenting after flowering; pubescent pubescent to glabrous leaves without lateral segments .…・.... 7a Achenes ciliate ..F. Xpurpurea

8a 8a Sepals inside glabrous ，outside curly pu 目 7b Achenes 紅 e not ciliate bescent ，usually green; pedicels and inflo- 12a Hypanthium at fruiting remains con- rescence rescence branches densely tomentose; cave ，ca 叩ophore absent; achenes flat ， leaves leaves beneath tomentose ， at least along rarely slightly twisted; leaves never veins; veins; stipules semiorbicular ，auriculate; tomentose beneath

petals petals with claws ，white. …...・ H ・..F. vestita 13a Petals pink; always hermaph- 8b 8b Sepals glabrous from both sides; inflores- rodite; stipules ovate to oblong-lan- cence cence entirely glabrous; leaves never ceolate ， not auriculate ，membranaceous tomentose tomentose beneath; stipules oblong-lanceo- 14a Petals with unevenly toothed margins late ，membranaceous ， not auriculate; pet- and reduced claws; pedicels and inflo- 306 306 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

rescence rescence branches glabrous; achenes 17b Plants usually less than 1m high ， glabrous ， with distinct stipes ，(2)3- generative shoots usually single

4( 4( 5) per flower .…...・ H ・...F. multijuga from rootstock ， with rosette; petals 14b 14b Petals with distinct claws and even less than 4 mm long ・… F. stepposa

m 訂 gins; inflorescence branches sp 訂 se shortly shortly bristly pubescent; achenes gla- A census of the Filipendula species brous ，sessile ，(1 )2-3(4) per flower.. Filipendula Mill. ， Gard. Dic t. Abr. ed. 4，512 (1754) ...... F.formosa ...... F.formosa - Adans. ，Fam. Pl. 2: 295 (1 763) - Maxim. in Acta 13b 13b Petals white; flowers pistillate ， Horti Petropo l. 6: 245 (1 879). staminate staminate or hermaphrodite; stipules Spiraea L.， Sp. Pl. ed 1，490 (1753) ，p.p. semiorbicular semiorbicular to ovate ， not mem- Ulmaria Hill ， Hort. Kew. ，213 (1 768). branaceous; branaceous; inflorescence branches Thecanisia Ra f.， New Fl. 2: 38 (1 837).

sp 訂 se tomentose Perennial rhizomatous herbs. Leaves interrupt- 15a 15a Flowers pistillate or staminate ，plants edly pinnatisected ， with palmately lobed terminalleaf

dioecious; dioecious; achenes 4-5 ， slightly segments. Inflorescence an anthela ，fruits apoc 紅 pous ， twisted ， stigma capitate …… achene-like ， with two ovules in each carpe l. … ..F. tsuguwoi Type: F. vulgaris Moench. 15b 15b Fl owers pistillate ，staminate or her- I. Sec t. Hypogyna (Shimizu) Schanzer ，sta t. nov. maphrodite ， plants polygamous; -ShimizuinJ. Fac. Tex t. Sc i. Techno l. Shinshu Univ. achenes achenes 5-9 ，flat; stigma decurrent . 10 ， se r. A ， 26: 4 (1 961) ， pro subgen.

…...・ H ・. F. kiraishiensis Hypanthiumnearlyflat ，sepals triangular ，toothed ， 12b 12b Hypanthium at fruiting becomes flat to with midribs; petals white ，clawless ， with a broad convex;ca 叩ophore more or less devel- base; achenes hispidly ciliate along sutures and on oped; oped; achenes twisted; leaves often both sides. Only one species. tomentose tomentose beneath Type: F. occidentalis (S. Wats.) Howell. 16a 16a Carpels and achenes glabrous ，much 1. Filipendula occidentalis (S. Wats.) Howell ， F l. twisted; twisted; pedicels and inflorescence NW Ame r. 1: 185 (1898) - Shimizu in J. Fac. Tex t. branches branches without tomentose pubes- Sc i. Techno l. Shinshu Univ. 10 ， se r. A ， 26: 4 (1961) cence; cence; stems never cobwebby in up- - Cronq. et Hitchc. ， F l. Pacif. NW 211 (1973).

per per part ・・ H ・H ・..…...・ H ・-… ..F. ulmaria Spiraea occidentalis S. Wats. in Proc. Amer.Acad. 16b 16b Carpels and achenes sinuate pubes- 18: 192 (1 883).

cent cent at the upper ~art of ventral su- Type: Oregon. ByTraskRiver ，Tillamook ， July 11 ture ，slightly twisted; pedicels and 1882 ， T. Howell (Syntypes: LE ，MO ，WTU). inflorescence inflorescence branches dense white- Distribution: USA ，NW Oregon. tomentose; tomentose; stems usually cobwebby Habita t: on rocks along mountain streams.

III III upper p紅 t Selected examined. specimens USA ，Or egon ，Tillamook Co. ，TraskRiver ，18.06.1936 ， Thompson 12750 ，topotype(MO ， 17 17 a Plants more than 1m high ，genera- US ，WTU); Tillamook Co. ， Wilson River ，23.07.1927 ， tive tive shoots:!::numerous from a root- Thompson 3136 (WTU); Tillamook Co. ， Trask River ， 29.05.1958 ，Hitchcock & Muhlick 21577 (WTU); Clatsop Co. ， stock ， without rosette; petals 4-5 Onion Onion Peak ，21. 07.1971 ，Chambers 3201 (WTU ，US).

mm long ...・ H ・H ・H ・.F. megaloc α rpa October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 307

Filipendula Filipendula occidentalis is a na 町 ow endemic and (1909) - Shimizu op. ci t. 6 (1961) ー Ohwi ，l. c. (1 965). possibly possibly a relic of the ancestral to the rest of the genus F. kamtschatica (Pall.) Maxim. f. glabra Koidz. group. group. lι(1909) - Ohwi ，1心(1 965). II. II. Sec t. Schalameya Juz. ， F l. URSS 10: 281 Type: Camtschatca ， . Pallas (Holotype: LE). (1 941) ， descr. ross.; Not. Sys t. (Leningrad) 17: 239 Distribution: Kamchatka ，Bering and Kuril isles ， (1955) ，emend. Schanzer hoc loco - Shimizu in Journ. Sakhalin ，Hokkaido ，N part of Honshu ，and two sites Fac. Fac. Tex t. Sci. Techno l. Shinshu Univ. 10 ，Se r. A ， 26: at the Okhotskoe Sea coas t. 5 (1961) ， p.p. Habita t: along mountain creeks and rivers ，on wet Folia Folia subtus haud tomentoso-pilosa ，segmentum slopes. Often grows in monodominant stands in river terminale terminale (3)5 (7)-palmatilobatum ，segmenta lateralia valleys of Kamchatka and Sakhalin. Filipendula lanceolata ，hypanthium post anthesin concavum camtsch αtica is the main component of so called ‘tall pe ロnanens. herb communities' of Kamchatka and Sakhalin com- Leaves lack tomentose pubescence ，a terminalleaf posed also of Cacalia hastata L.， Senecio cannabifoli ω segment (3)5(7)-left ， lateral ones lanceolate ，a Less. ， Urtica pl αη Iphylla Wedd. and a few other hypanthium after flowering remains concave. The specles. section section is delimited mostly by overlapping characters Selected Selected specimens examined. Russia ，Kamchatka ， mouth of the species and series. ofKichigaRiv. ，Smetanin(VLA);Karaginskyiis l.， 22.08.1976 ， Gorshkov Gorshkov (VLA); footofShiveluch Volc. ，13.08 .1 978 ，Sofeikova A. Ser. Schalameyae ， se r. nov. (MHA); (MHA); W coast ，Oblukovina Riv. ，23.08.1936 ， Liubimova Petala Petala alba ， sepala intus appresse pilosa ， achenia (LE); Shchapino ，3.08.1909 ，Komarov (LE); Chazhma Cape at E coast ，19.08.1920 ， Skobunov (LE); Bering is l.， 6.09.1929 ， ciliata. ciliata. Kardanova Kardanova 202 (LE). Kuriles ， Onekotan is l.， 13.08.1962 ， Petals Petals white ，sepals appressed-hairy inside ， achenes Cherniaeva (LE); Shumshu is l.， 8.08.1955 ， Mishin (LE); Shiashkotan Shiashkotan is l.， 25.09.1956 ，Stepanova (VLA); Iturup is l.， ciliate. ciliate. 29.08.1961 ， Woroschilow 10830 (MHA); Kunashir is l.， Type: type of the section. 25.09.1987 ，Schanzer (MHA). Sakhalin ，N part ， Nyljvo Riv. ， 7.09.1908 ，Komarov (LE); Tymovskoe ，17.09.1962 ， 2. 2. Filipendula camtschatica (Pall.) Maxim. in Acta Woroschilow Woroschilow 11472 (MHA); Alexandrovskyi ，19.07.1906 ， Horti Horti Petropo l. 6: 248 (1 879) (Maxim. scripsit Shestunov 217 (LE); Nevelsk Distr. ，Kuznetzovo ，6.07.1950 ， Kravchenko Kravchenko (LE). N coast of Okhotskoe Sea ，Piyagin penins. ， ‘kα mtsch αtica'). Yakubov Yakubov (VLA). Amur mouth ，Orelj Lake ，19.08.1934 ヲ Zinger F. F. kamtschatica auc t.: Juz. ， F l. URSS 10: 281 (LE). Japan ，Hokkaido ，Oizumi ，30.07.1980 ，Boufford & Kato 22488 (GH); Sapporo ，5.07.1903 ， Arimoto (GH); Honshu ， (1941) (1941) -Shimizu inJ. Fac. Tex t. Sci. Techno l. Shinshu Nikko ，Tochigi Pref. ， Aug. 1930 ， Makino (MAK); Akita Pref. ， Univ. Univ. 10 ， Ser. A ， 26: 5 (1 961)) - Ohwi ， F l. Jap. ，537 Kazuno-gun near Hakka-toge ，15.07.1971 ，Naito & al. (MO); Iwate Iwate Pref. ， M t. G 弔問， 28.08.1894 ， Faurie 13676 (MO); (1 965) - Worosch. ， Opredelitel Ras t. Soviet. Daln. Yamagata Yamagata Pref. ， M t. Chokai-san ，17.08.1960 ， Ohashi 6728 Vos t.， 354 (1 982). (GH). Spir α ea camtschatica Pall. ，F l. Ross.l: 41 (1784). The Koidzumi' s forms are not recognized herein S. S. palmata auc t. non Pall.: Thunb. ， F l. Jap. ，212 because the variety of leaf and stem pubescence is (1 784) ， p.p. much more diverse. It seems not reasonable to de- Ulmaria Ulmaria kamtschatica (Pall.) Yabe et Yendo in scribe it by taxonomic means. U sually minute Bo t. Mag. Tokyo 28: 186 (1 904). hispidulous pubescence on pedicels and inflorescence Filipendula Filipendula kα mtschatica (Pall.) Maxim. f. typic α branches is very characteristical of F. camtschatica ，

Koidz. Koidz. inBo t. Mag. Toky023: 108 (1 909)-Ohwi ，1心 though it is sometimes absent in the southern part of (1965). (1965). its range. Stems and inflorescence branches can be F. F. kamtschatica (Pall.) Maxim. f. pilosa Koidz. l. c. glabrous to densely hispid ¥with rough long hairs. 308 308 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

Stem pubescence usually does not correlate with that Nakai ，n.v.; Koang-neug ，1912 ，Mori 30 ，259 ，n.v. (a of of leaves and petioles. specimen with such label but without number Shimizu 3. 3. N akai ，Feddes Reper t. 13: (1 961) cited as the lectotype); in herbidis 274 (1914) - Shimizu in J. Fac. Tex t. Sci. Techno l. Chirubulsusan ，9.07.1913 ，Nakai 695 ， v. photo.; in Shinshu Shinshu Univ. 10 ， Ser. A ， 26: 7 (1961) - Worosch. ， herbidis m t. Chandorobon ，1400 m et supra ，6.07.1913 ， Opredelitel Opredelitel Ras t. Sovie t. Daln. Vos t. 354 (1 982). Nakai ， in n.v.; herbidis m t. Chirisan ，08.1912 ，Mori F. F. koreana Nakai ， Fl. Sylv. Kor. 7: 8 (1918) ，nom. 168 ， n.v.; ibidem. ，30.06.1913 ，Nakai 68 ， n.v. nud. nud. (Syntypes: TI).

F. F. ciliat α(Koidz.)， Kudo ，Veg. Yezo 263 (1 925) ， Distribution: Primorsky and Khabarovsk Prov- p.p. p.p. inces of the Russian Far East ，NE China ，Korea ， F. F. glabra N akai ex Kom. et Aliss. ，Key Pl. Far East Hokkaido and Kunashir Islands. Reg. Reg. URSS ，2: 653 (1 932) ，nom. nud.-Juz. ，Fl. URSS Habita t: banks of mountain creeks in forests and 10: 10: 281 (1 941). meadows ， usually in small groups. In Kunashir (and F. F. yezoensis Hara ，J. Jap. Bo t. 10: 235 (1 934) - Hokkaido?) grows usually in meadows and scrub

Shimizu l. c. (1 961) - Worosch. ， l. c. (1 982). instead of s佐eam banks occupied by communities of F. F. purpurea auc t. non Maxim.: Kom. et Aliss. ， op. F.c αmtschatica. t.， ci t.， 650 (1 932) - Juz. ， op. ci t.， 282 (1 941) - Li in Yu Selected examined. specimens Khabarovsk Prov. ， Mt.

Tardoki 聞 Yani ，29.08.1981 ， Gotvansky 6059 (MHA); Ak ur et et al.， Fl. Reip. Pop. Sin. 37: 7 (1 985). Valley Valley (tributary of Tumnin Riv よ25.07.1972 ， Nechaev 133 F. F. multijuga Maxim. var. ciliata Koidz. in Bo t. (お 1HA); BikinDis 江， ZhivkaRiv. ，11. 08.1980 ，Schlotgauer398 (MHA); (MHA); Pri morsky Pr ov. ，OlgaBay ，7.07. 1955 ，Egorova(1 位IA); Mag. Tokyo 23: 178 (1909) ，quoad pl. ex Hokkaido. S Sikhote-Ali 吋， M t. Labalaza ，15.09.1930 ，Schischkin 1094 F. F. multijug αMaxim. var. koreana Nakai ，1ι (LE); Khanka Distr. ，Kamenj-Rybolov ，28.06 .1 928 ，Vorobiev (LE); (LE); Tumannyi Cape ，15.07.1944 ， Zhudov 2098 (MW); (1 914). Vladivostok ，28.07.1985 ， Schanzer (MHA); Kunashir ， F. F. multijuga Maxim. v紅 . alba N akai ，1ι(1914). Golovnino ，24.09.1987 ，Schanzer (MHA). China. Liaoning Prov.: Prov.: Fusan distr. ，Mangtzyang ，28.07.1950 ，Wang Sui-Kang F. F. multijuga Maxim. va r. yezoensis Hara ，1ι 1810 1810 (LE); Heiloungkiang Prov. ，Shangtze Dist r.， Weihe ， (1 934) ， pro syn. - Ohwi ， F l. Jap. ，537 (1 965). 19.08.1953 ，T. H. Lion & Baranov 1386 (LE). Korea. Kangwon Prov. ， M t. Hyangnobong ，18.06.1967 ，Y. H. Chung (GH); Hogen F. F. multijuga Maxim. subsp. yezoensis (Hara) Prov. ，Hongo-san ，7.07.1918 ， Wilson (US ， GH); J apan ， Woros ，C h. ，Bull. Soc. Nat. Moscou 96: 127 (1991). Hokkaido: M t. Yubari ，3.08.1965 ，Shimizu (SHIN); M t. Apoi ， 20.08.1966 ，Shimizu (SHIN). F. F. multijuga Maxim. subsp. koreana (Nakai) The and nomenclature of F. glaberrima Worosch. ，1ι (1 982) - id. ，l. c. (1 991). remains remains rather confusing till now. The species has F. F. kamtsch αtica (Pall.) Maxim. v訂 . glaberrima been described several times by different authors Nakai in Bo t. Mag. Tokyo 26: 128 (1 913) ，nom. nud. under under different names from different parts of its F.c αmtschatica (Pall.) Maxim. subsp. glaberrima range. range. The identity of some of these taxa has been (Nakai) (Nakai) Worosch. ， op. ci t. 126 (1 991). already already discussed in literature (Woroschilow 1960 ， F. F. yezoensis Hara f. alba (N akai) Shimizu ， op. ci t.， Shimizu 1961). FirstNakai (1914) describedmeadow 9 (1 961). and forest forms of this taxon as separate taxa ， F. F. F. yezoensis Hara var. hispida Shimizu ，1ι (1 961). glaberrima glaberrima and F. multijuga v訂 . koreana ，respec- Spiraea Spiraea digitata Willd. var. galbra Ledeb. ， F l. tively. tively. Later on he produced the nomen nudum F. Ross.2: Ross.2: 18 (1 844). koreana ，which Shimizu (1 961) synonimized with F. Type: Type: Corea media et australe ， M t. Namahansan ， yezoensis yezoensis Hara from Hokkaido. 1 have not seen the 2.08 .1 902 ， T. Uchiyama ，n.v.; Quensan 9.06.1909 ， October October 1994 Journal of Japanese Botany Vo l. 69 No. 5 309

type type of F. yezoensis to compare it with that of F. Filipendula camtschatica ， F. glaberrima and F.

multijuga multijuga v紅 . koreana ， as Shimizu (1961) has done ， multijuga 訂 e quite distinct though related species ，

but but the materials I have seen from Hokkaido and well sep 訂 able from each othe r. Kunashir Kunashir Islands are really not separable taxonomicall y B. Ser. Formosae ， se r. nov. from those from Korea and Promorsky Prov. of Rus- Achenia glabra aut raro ciliata (tantummodo F. sia ，including the isotype of F. multijuga var. kore αna multijug αvar. ciliata) ，stipitata aut sessilia ， petala (North (North Korea ， Ibaien pass near Naran ， Jalu Riv. ， rosea ，stipulae ovata aut oblongae ，membranaceae ， 6.07 .1 897 ， V. Komarov ， LE). In the same paper haud auriculatae. Shimizu Shimizu (1961) found out that F. purpurea sensu Achenes glabrous ，rarely ciliate (in F. multijuga Komarov and Alissova (1 932) and Juzepczuk (1941) ciliata) var. ，stipitate or sessile ，petals pink ，stipules is is by no means the J apanese F. purpurea Maxim. ， but ovate to oblong ，membranaceous ， not auriculate. quite quite a separate species ，which he also identified as F. Type: F.formosa Nakai. yezoenSls. yezoenSls. 4. Filipendula formosa Nakai in Feddes Reper t. 13: Filipendula Filipendula glabra (Maxim.) Nakai is another 274 (1 914) - Shimizu in J. Fac. Tex t. Sc i. Techno l. confusing confusing name created by Komarov and Klobukova- Shinshu Univ. 10 ，Ser. A ， 26: 19 (1 961). Alissova Alissova (1932) evidently by mistake ，and never Type: Corea. In herbidis declivitatis montis Sin- described described by Nakai nor Maximovicz. This fact was Yang-chini ，finis occidentalis Chirisan ，70 0- 900 m ， discovered discovered by Woroschilow (1960) who synonimized 13.07.1913 ， T. Nakai (Holotype: TI ，v. photo). the the latter species withF. koreana ，i.e. with F. yezoensis Distribution: an endemic of South Korea known sensu sensu Shimizu (1961). On the other hand ， Shimizu only from M t. Chirisan. (1961) (1961) supposed thatF. g labra is no doubt synonimous Habitat: meadows. with with F. glaberrima. The identity of the remained F. Specimens examined. Korea ，Chirisan ， South Chulla ， glaberrim αandF. yezoensis was shown by Schanzer 9.08 .1 934 ，Smith (GH); ibid. ，15.08.1934 ，Smith (US). (1989c). (1989c). Theonlydifferenceof F. glaberrimafromF. Filipendula formosa is closely related to F. yezoensis yezoensis supposed to be the presence of 1-4 shortly multijuga ， a vicarious species of Japan ， bearing iden- stipitate stipitate to sessile achenes instead of 5-81ong stipitate tical membranaceous stipules ， pink petals and gla- brous brous achenes. Still it resembles F. camtsch αtica in a ones. ones. It appe 訂 ed that these differences fall into the variability variability amplitude of one species and can be found number of characters (hispid pubescence ，leaves with within within a single population. a small number or without lateral segments ， sepals Woroschilow (1 982) distinguishedF. glaberrima appressedly pubescent inside ，petals with claws) and from F. koreana by its larger overall dimensions ， seems to be intermediate between them to some reniform ， never glaucous beneath lobes of terminal exten t. leaf leaf segments ，and larger infloresences ，which do 5. Maxim. in Acta Horti nothing nothing with the diagnostic characters of F. glaberrima. Petropo l. 6: 247 (1879)- Shimizu inJ. Fac. Tex t. Sci. All All the above characters are very variable due to Techno l. Shinshu Univ. 10 ， Ser. A ， 26: 11 (1961) - environmental environmental conditions. I can also hardly agree Ohwi ， F l. Jap. ，527 (1 965). with with his recent suggestion (W oroschilow 1991) to F. multijuga Maxim. v訂 .ciliata Koidz. ，Bo t. Mag. regard regard F. glaberrima as a subspecies of F. Tokyo23: 179 (1 909) ，quoad pl. exHonshu-Shimizu camtschatica ， as well as F. koreana andF. yezoensis ， op. ci t. 12 (1 961) - Ohwi ， l. c. (1965). treated treated as separate taxa ， as subspecies of F. multijuga. Type: Japonia ，Jokohama ，Hakone in cacumine 310 310 植物研究雑誌第69 巻第5 号 平成 6 年 10 月 montium lapidosorum graminosorum ，18-30.10.1862 ， that possess ciliate achenes and appressedly pubes- Maximowicz (Lectotype: LE). Nagasaki ，Kundsho- cent inside instead of glabrous sepals. Both characters san san ad pedem ，28.09-9.10.1863 ，Maximowicz; in can be regarded either as an example of homologous montibus montibus tractus Aidzu ，08.1877 ， Faurie (Syntypes: variability ，bringing to light some traits typical for the LE). LE). series Schalameyae ， or the result of an introgression Distribution: Distribution: Japan ，Kyushu ，Shikoku and south of from F. camtschatica. The latter has its southem Honshu Islands. distributionallimit in middle Honshu where hybrid- Habita t: mountain forests and subalpine mead- izes with F. multijuga ， as has been described above. ows. ows. C. Ser. Dioeciae ， se r. nov. Plantae Plantae dioeciae aut polygamae; achenia glabra ， Se1ected Se1ected specimens examined. Japan ， M t. Kuju ，Oita Pref. ， Makino Makino (MAK); Ashigaroshimo-gun ，Kanagawa Pref. ， sessilia aut brevissime stipitata ， petala alba; ramuli 14.07.1938 ， Makino 129335 (LE); Oizumi ，Nerima-ku ， Tokyo inflorescentiae tenuiter tomentos i. Pref. ，24.06.1939 ， Makino (LE); Taira ，Omachi-city ， Nagano Pref. ，1927 ，お 1akino 129350 (LE); M t. Mitsutoge ，Yamanashi Dioecious or polygamous plants; achenes gla- Pref. ，1930 ， Makino (LE); Japan A1ps ， 40 mi. of Toyama ， brous ，sessile or very shortly stipitate ，petals white; Charette Charette (MO); Hira Mts. ，Shiga Pref. ，8.08.1980 ，Boufford & Kato Kato 22677 (MO ， GH); Mt. Daifugen-dake ， Nara Pref. (GH); inflorescence branches sparse tomentose. M t. IwaguroinIshizuchiMts. ，EhimePref. ，18.07. 1982 ，Murata Type: F. kiraishiensis Hayata. & al. 190 (GH); M t. Gozaisho ，Ise Prov. ，Kitamura (US). 6. 6. Filipendula kiraishiensis Hayata ，Ic. Pl. Formos. Filipendula Filipendula multijuga is a rather variable species. 9: 9: 39 (1 920) - Shimizu in J. Fac. Tex t. Sci. Techno l. It s subalpine dwarf ecotype (to which all the type ShinshuUniv.l0 ，Ser.A ， 26: 18(1961)-Suin Li net materials materials belong) is strikingly different in habit from l.， al.， Fl. Taiw. 3: 67 (1 977) - Li in Yu et al.， Fl. Reip. the the forest one. Still there is no difference between Pop. Pop. Sin. 37: 7 (1985). them in taxonomically meaningful characters. Plants Type: Type: Kiraishi ，ad 10 ，000 ped. al t.， 08.1918 ， S. with with white instead of pink flowers also occur haphaz- Ohashi (Holotype: TAIF ，n.v.). The type specimen is ardly ardly throughout F. multijuga range. Plants with cili- destroyed destroyed representing a single leaf only (Shimizu ate ate instead of glabrous achenes sometimes occur at 1961) 1961) the the northem border of F. multijuga range in central Distribution: Distribution: central and northem Taiwan. Honshu. Th e latter were described by Koidzumi (1909) Habita t: rocks in subalpine zone at elevations ca. as as var. -ciliat α，which was originally reported from 3，000-3 ，700 m. 'J 'J aponia media et septentrionali' without quoting any Specimens Specimens examined. Taiwan ，M t. Kiraisyu-nampo ， specimens. specimens. Several years after Koidzumi (1 913) indi- 24.08.1929 ，Suzuki 2408 (MO); Nantou Co. ， M t. Hohuanshan ， cated cated a number of specimens of this variety from 2.08.1986 ，C. 1. Peng 9595 (恥 1HA); ibid. ，13.07.1985 ，C. I. Peng 8266 8266 (MHA); I1 an Co. ，Nan-hu-bei-shan ，16.08.1983 ，C. I. Peng central central and northem Honshu and Hokkaido. How- 5792 (MHA). ever ， those from Hokkaido appeared to belong to F. Shimizu (1961) revealed that this species is po- g laberrima (Shimizu 1961). As for those from Honshu ， lygamous. The materials 1 have seen confirm this they they all but one (Shimizu 1961) tumed to be los t. Th e opinion. opinion. Decurrent stigmas are one more peculi 訂 latter latter specimen designated by Shimizu as the Lectotype character of this species ，which 1 believe to be highly (Nikko ，07.1885 ，Matsumura ，TI ，v. photo) unfortu- plesiomorphic. On the other hand ， F. kiraishiensis nately nately bears only buds and no mature achenes. Still 1 resembles F. multijuga in leaf shape and pubescence ， have been lucky to study two specimens (Oizumi ， and to some extent in the stipules and achenes shape. Nerima-ku ，Tokyo Pref. ，20.07.1935 ，Makino ，MHA) 7. Filipendula tsuguwoi Ohwi ，Acta Phytotax. October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 311

Geobo t. 15: 115 (1 954); Fl. Jap. ，537 (1965)-Shimizu Univ. 10 ， Ser. A ， 26: 13 (1961) - Li in Y 品目 al.， Fl. inJ. inJ. Fac. Tex t. Sci. Techno l. Shinshu Univ. 10 ，Se r. A ， Reip. Pop. Sin. 37: 6 (1 985) - Worosch. ，Opredelitel 26: 26: 19. Rast. Sovie t. Daln. Vost. ，354 (1 982).

Type: Type: M t. Ishitate in Iyo ，1700 m ，Yamanaka Filipendula palmata (Pall.) Maxim. v訂 .tomentosa 10913 (Holotype: TNS ，n.v.). (Ledeb.) Kom. ， F l. Mansh. 2: 522 (1904). Distribution: Distribution: Kyushu and Shikoku Islands. F. rufinervis Nakai in Bo t. Mag. Tokyo 26: 35 Habita t: gravelly places of limestone or conglom- (1912). erate erate in high mountains over 1000m (Shimizu 1961). F. nuda Grub. ，No t. Sys t. (Leningrad) 12: 112 1 have failed to get any specimen of F. tsuguwoi (1950). and rely upon descriptions and figures in Ohwi (1954 ， F. amurensis A. Baran. in Bo t. Jahrb. 79: 523 1965) 1965) and Shimizu (1 961). Filipendula tsuguwoi is a (1960). dioecious dioecious species mo 叩hologically strongly resem- F. palmata (Pall.) Maxim. f. nuda (Grub.) Shimizu blingF. blingF. kiraishiensis. It s distributionhas beenmapped op. ci t. 14 (1961). according according to specimens quoted in Shimizu (1961). Spiraea palm α ta Pall. ， Reise Prov. Russ. Reichs 3: Holotype Holotype is represented by a staminate plant ，paratypes 735 (1 776). are are pistillate: 'M t. Komochigongenyama in Tosa ，ca. S. digitata Willd. ， Sp. Pl. 2: 1061 (1 779).

1600 m ，Yamanaka 7307; Mt. Ishizuchi in Iyo ， Spiraea digitat αWilld. v訂 . tomen ω sa Ledeb. ， F l. 25.07.1947 ，Morioka 80187 (TNS ，n.v.)'. Ross. 2: 17 (1 844).

1. 11 1. Sec t. Albicoma Juz. ， Fl. URSS 10: 283 ，descr. S. digitata Willd. vぽ . latiloba Glehn ，Acta Horti ross.; ross.; No t. Sys t. (Leningrad) 17: 240 (1 955) ，emend. Petropo l. 4: 37 (1 876). Schanzer Schanzer hoc loco - Shimizu in J. Fac. Tex t. Sci. Ulmaria palmata (Pall.) Focke in Eng l. et Prantl ，

Techno l. Shinshu Univ. 10 ， Ser. A ， 26: 15. N 剖. Pfl. Fam. 3， 3: 41 (1894). Plantae Plantae rhizomate longis stoloniformibus; folii Type: In regionibus alpinis transbaicalensibus... segmentum terminale profunde 7-11 palmatilobatum ， (The locality ofthe type specimen is unknown). 1 have segmenta lateralia 2- 510bata; achenia elongata ， breve failed finding the type specimen in LE ，it also was not stipitata stipitata aut sessilia. found in K and BM through my reques t. Plants Plants with long subterranean runners; terminal Distribution: E Siberia ，NE Mongolia ， Russian Far leaf leaf segment deeply palmately cleft into 7-11 lobes ， East ，NE China and N Korea. lateral lateral leaf segments cleft into 2-5 lobes; achenes Habita t: wet sites along rivers ，forests ，meadows ， elongate ， with short stipes or sessile. roadsides. Type: Type: F. angustiloba (Turcz.) Maxim. Selected specimens examined. Russia ， E. Siberia ， Krasnoyarsk Krasnoyarsk Prov. ，Igarka ，31.08.1947 ，Tyrzhikov (MW); 5 km D. Ser. Albicomae. W of Enisseisk ，17β7.1947 ， Evseeva (LE); Baikal Lake ， Folia Folia subtus tomentosa vel solum ad nervos pilosa ， Listvennichnoe ，3.08.1955 ，Sokolov 296 (LE); Podkamennaya Tunguska Tunguska Riv. ，29.07.1909 ，Borovikov (LE); Vitmin Riv. ， petala petala alba ， sepala intus glabra. 8.07.1911 ，Sukachev (LE); Tchita ，10.06.1912 ，Krasnopolskaya Leaves usually tomentose beneath ， petals white (LE); Yakutia ，Khalamandy Riv. (right tributary of Lena Riv よ 30.07 .1 953 Kuvaev (MW); NE Mongolia ， 120 kmE ofDarkhan ， with with even margins ， sepals inside glabrous. 16.07.1981 ， Gubanov 3471 (MW); Russian Far Eas t: Bl agowesh- Type: Type: type of the section. chensk ，2.07.1891 ，Korzhinsky (LE); Amur-Bur 吋a lowland ， Pojarkowo ，1.07.1914 ，Kr ischtofowicz 301 (LE); Amur Prov. ， 8. 8. Filipendula palmata (Pall.) Maxim. in Acta Horti Tukuringra Tukuringra range ，3.08.1971 ，Golysheva (MW); Lower Amur ， Petropo l. 6: 250 (1879) - Juz. ， F l. URSS 10: 282 Khungari Riv. ，06.1930 ，Kutyseva (LE); Okhotsko-Kolymsky Prov. ，Tauysky distr. ，09.1955 ，Antonova (LE); 恥1agadan Prov. ， (1 941)-ShimizuinJ.Fac. Text. Sci. Techno l. Shinshu 312 312 植物研究雑誌第69 巻第5 号 平成 6 年 10 月

Ann anj ，26.08.1969 ，Khokhryakov (MHA); Kamchatka ，Tigilj ， Distribution: NE China and the contiguous areas 9.08 ，1960 ， Baskin (MW); Kamchatka ，Elizovo ，12.08.1960 ， Woroschilow9767 Woroschilow9767 (MHA); Sakhalin ，Tymovskoe ，12.08 .1 955 ， of Russia and Mongolia. Shukhobodsky Shukhobodsky 230 (LE); Primorsky Prov. ， Tetyukhe Riv. ， Habita t: wet meadows. 18.06.1964 ，Chizhevskaya (MHA); Vladivostok ，10.09 .1 950 ， Selected Selected specimens examined. NE China ，Inner Mongolia ， Woroschilow Woroschilow 4867 (MHA). NE China ，In ner Mongolia ，Huna near near Hailar M t.， S. H. Li et al. 1126 (LE); Dzalatun ， Great Prov 円 Argunj Dist r.， 16.08.1951 ， S. H. Li 1978 (LE); Khingan ，27.07.1938 ，B. Skvortzov (GH); Harbin ，29.06.1937 ， Heilungkiang Heilungkiang Prov. ，Great Khingan ，Y ak -shan range ， 7 .07 .1909 ， Ivashkevich Ivashkevich 171 (LE); Liaoning Prov. ，Jingyui Distr. ，2.07 .1 950 ， B. Skvortzov (GH); N Manshuria ， Mergen to Aigun ，13.06- 13.07.1910 ， Ladygin (LE); Kirin Distr. ，Ninguta ，5.07.1923 ， T. T. N. Liou 1205 (LE) ‘ Corea ，3.07.1956 ， Pong Syup Joh 174 (LE). (LE). Tretiakoff (LE). Mongolia ， East Aimak ，W parts o~ Great Khingan ，17.07.1985 ， Gubanov 9880 (MW). Russia ，Urulungui Usually Usually leaf lobes of F. palmat αare oblanceolate Riv. near Klichka ， 1. 08.1930 ， Ivanova 243 (LE); Bl agoweshchensk ，20.06.1904 ， Karo 216 (LE); Amur Valley and deeply double se 町 ate. Plants with lanceolate between between Khabarovsk and Troitzkoe ，15.07.1926 ， Neimark 64 weakly se 町 ate leaf lobes ， occurring throughout its (LE); Vladivostok Dist r.， W shore ofKhanka lake ，2.08.1926 ， range ，were described as F. αmurensis. Leaves of F. Melvill (LE). palm α ta are almost always grey tomentose beneath ， at 1n contrast to F. palmat α，F. αngustilob αis vari 国 least least the uppermost ones. Plants that lack tomentose able as to leaf pubescence: both forms with tomentose pubescence are extremely rare and known as F. nud α and appressedly pubescent beneath along veins leaves (e.g. ，‘ N erczinsk' Transbaikal area ，06.1899 ，Mauritz ， occur simultaneously. Specimens with) tomentose holotype ， LE); Transbaikalia ，Karymskaya ，3.08.1928 ， leaves Juzepczuk (1 941) called F. intermedi αf. leiocarp α. 1 treat them as F. αngustiloba ，because they Solokhin Solokhin 110 ，LE; Yakutsk Prov. ，Sp 紅 t of 01 凸kma Dist r.， Tungir Riv. ，28.07.1910 ，Sukaczev 1250 ， LE). possess all the other diagnostic characters ofthe latter. Both forms do not deserve the specific rank. E. Se r. Rubrae E. Sergievskaya emend. 9. 9. Filipendula angustiloba (Turcz.) Maxim. in Acta Folia subtus haud tomentosa; achenia breve HortiPetropo l. 6: 250 (1 879)-Juz. ，Fl. URSS 10: 288 stipitata ，glabra; petala rosea ，margine inaequaliter (1941)-ShimizuinJ.Fac. Tex t. Sci. Techno l. Shinshu dentata; sepala intus appresse pilosa. Univ.10 ，ser. A ， 26: 15 (1 961)- Worosch. ，Opredelitel Leaves lack tomentose pubescence; achenes shortly Ras t. Sovie t. Daln. Vos t.， 353 (1 982) - Li in Yu et al.， stipitate ，g1abrous; petals pink ， with uneven1y toothed l. F l. Reip. Pop. Sin. 37: 9 (1 985). margins ， sepa1s appressed1y pubescent inside. F. F. intermedia (Glehn) Juz. ， op. ci t.， 284 (1 941) ， Type: F. rubra (H il1.) Robins. p.p. ，quoad f. leioc α rpa Juz. ，desc r. ross.; No t. Syst. 10. Filipendula rubra (H il1) Robins. in Rhodora 8: 204 (1 906) - Shimizu in J. Fac. Tex t. Sci. Techno 1. (Leningrad) (Leningrad) 17: 240 (1 955) ー Li in Yu et al.， op. ci t.， 9 (1985). Shinshu Univ.l0 ，Ser. A ， 26: 14 (1961)-G1eas. ，New Spir αωα ngustiloba Turcz. in Fisch. et Mey. ， 1nd. Brit t. & B r. Fl. North U.S. & Canad. 3: 299 (1963). Sem. V1U Hor t. Bo t. Petropo l. 71 (1 841). Filipendula lob α ta (Gronov. ex Jacq.) Maxim. in S. S. lobata Murr. var. angustiloba Turcz. ， F l. Baic.- Acta Horti Petropo 1. 6: 251 (1 879). Dahu r. 1: 364 (1 843). Spiraea rubra (Hi l1) Brit t.， Bu l1. Torrey Bo t. Club S. S. digitata Willd. var. angustiloba (Turcz.) Glehn ， 18: 270 (1 891). Acta Horti Petropo 1. 4: 38 (1876). Thecanisia lobata (Gronov. ex Jacq.) Raf. ，New S. S. digitata W iI1 d. var. intermedia Glehn ，1心(1 876) ， Fl. North Amer. 2: 38 (1 836). p.p. p.p. T. ponpurea Raf. ，l. c. (1836). Type: Type: 1n pratis Dahuriae ，Nerczinscoi Zavod ，1831 ， T. angust 扮lia Raf. ，l. c. (1836). Turczaninov. Turczaninov. (Lectotype and isolectotypes: LE). UlmariarubraHill ，Hor t. Kew. ，214 ，tab. 7 (1 768). October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 313

Type: Type: Ex America boreali. (The locality of the type F. vestita (Wal l. ex Hoo k. f.) Maxim. f. viridula specimen specimen is unknown). Purchit et Panigrahi in Bangladesh J. Bo t. 10: 97 Distribution: Distribution: NE of the U.S.A. (1 981). Habita t: wet prairies ，fens ，rarely in forests; culti- Spiraea kamtschatica Pall. var. him αlensis Li nd l. vated vated as omamenta l. in Bo t. Reg. 17: 4 (1 841). S. S. vestita Wal l. ex Hoo k. f. depauperata Aitch. Selected Selected specimens examined. USA ，Michigan ，Emmet Co. ，Douglas Lake Region ，10.08.1924 ，Ehlers 3054 (LE); in J. Linn. Soc. 18: 52 (1880). N ova Scotia ，Yarmouth ，23.07.1921 ，Femald et al. 23965 (LE); S. vestita Wall. ex Hook. f. ， F l. Bri t. Ind. 2: 323 New Hampshire ，Sisbon ，27.07.1885 ，Safford 788 (US); Ill i- nois ，Beardstown ，07.1842 ， Geyer (LE); Conn. ，Hairfield ， (1 879). 11. 07.1893 ， Eames (US); CarolinaSep t.， adSawanno ，08.1842 ， S. ulmaria auc t. non L.: Boiss. ， F l. O r.， supp l.， 230 Rugel Rugel (US); Mass. ，Sheffield ，27.07.1920 ，Church i1l (MO); Indiana ， Miami Co. ，Peru ，1. 07.1942 ，Charles (WTU ， MO); (1 888) ，quoad pl. Aitch. Ohio ，Cincinnati ，14.07.1904 ，Felter (MO); Missouri ，St. Francois S. kamtschatica auc t. non Pall.: Wall. ，Ca t. n. 704. Co. ，24.06.1981 ，Orzell 321 (MO); NY ，Clifton ，1. 07.1916 ， Ruth Ruth (US); Virginia ，N ofM i1l boro ，18.07.1947 ，Freeman (US). Type: Ex alpibus Kumoon ， R. Bankworth IV. IV. Sec t. Filipendula. (Holotype: K ， cibacopy of the holotype: MHA). Hypanthium post anthesin convexum ， torus in Distribution: E Yunnan and Szechuan ，W Himalaya carpophorum breve elongatus aut inconspicuus; up to Afghanistan. achenia achenia breve stipitata aut sessilia ， ad latera carpophori Habita t: stream banks and wet slopes in the tem- affixa; affixa; achenia hilo laterali. perate mountain bel t. Hypanthium after flowering convex ， short Selected specimens examined. China ，Yunnan ，between Chungtien Chungtien Riv. and the Yangtze ，07.1923 ， Rock 9801 (US); carpophore carpophore usually developed; achenes shortly stipitate between Li kiang ， Y oungning and Youngpei (SW Szechuan) ， to to sessile ，laterally attached to torus. 05 --0 6.1922 ， Rock 5308 (US ， GH); in montis Hee-chan-men supra supra Hokin (Prov. de Yun-nan) ，Delavay 2525 (GH). Nepal ， Type: Type: type of the genus. near Tarakot ，Bheri Riv. ，13.07.1952 ， O. Polunin & al. 2436 F. F. Ser. Ulmariae (Mill.) Schanzer ，sta t. nov. ，mu t. (GH). India ，Kashmir ，Sind Valley ，23.06.1892 ，Duthie 11436 (LE); (LE); Punjab ，Rotang pass ，9.09.1935 ， Nath 389 (GH); W char. char. -M i1l.， Gard. Dic t.， Abr. ed. 4: 1490 (1754) ， pro Himalaya ，NW to Tsamba ，9-16.07.1856 ，Schlagintweit 3643 gen. gen. - Juz. ，Fl. URSS 10: 284 (1 941) ， pro subgen.- Pangi (GH); ，Bushahr ，Simla ，14.07.1934 ，Parmanand 510 (GH); (GH); Kumaon ，Starchey & Winterbottom (LE). Pakistan ，Bashar Shimizu Shimizu in J. Fac. Tex t. Sci. Techno l. Shinshu Univ. gol ，distr. Chitral ，17.06.1977 ，Muquarrab Shah & Dilawar 10 ，Ser. A ， 26: 20 ， pro sec t. (GH). Afghanistan ， Kurrum Valley ，1879 ，Aitchison (GH ，LE). Achenia Achenia matura plus minusve contorta; folia subtus Leaves of F. vestita can be tomentose beneath or tomentosa tomentosa vel solum ad nervos pilosa. pubescent only along veins (ι viridula). Some speci- Mature Mature achenes more or less twisted; leaves often mens ofYunnan evidently possess long subterranean tomentose tomentose beneath. runners: base of the E flank of the Lichiang Range ， Type: Type: F. ulmaria (L.)乱I{ axim. 09.1910 ，Forrest (US); Rock 9801 (US). This brings 11. 11. Filipendula vestita (Wall. ex Hoo k. f.) Maxim. in them close to F. palmata and F. angustiloba of Sec t. ActaHortiPetropo l. 6: 248 (1879)-Shimizu inJ. Fac. Albicoma which also possess tomentose leaves. It Tex t. Sci. Techno l.， Shinshu Univ. 10 ，Se r. A ， 26: 17 evidently implies common ancestry of both sections (1961) (1961) - Sch 凸nb.-Tem. in Rech. f. ， F l. Iran. 66: 122 or possible derivation of F. vestita from some extinct (1 969) - Li in Yu et al.， Fl. Reip. Pop. Sin. 37: 6 member of sec t. Albicoma. West Himalayan (1 985). populations of F. vestita ，however ， lack this character

Fili) フendula vestita (Wall. ex Hoo k. f.) Maxim. f. and more resemble F. megalocarpa ， remaining yet depauperata depauperata (Aitch.) Shimizu 1心(1 961). conspecific with the Yunnan populations. 314 植物研究雑誌第69 巻第5 号 平成6 年 10 月

12. 12. Filipendula megalocarpa Juz. in No t. Sys t. Type: Bashkir Republic ， steppe to the E of (Leningrad) (Leningrad) 17: 240 (1 955). Lomovtzev' s farm ，a sinking of microrelief ，Ar gayash F. F. ulmaria (L.) Maxim. f. megalocarp α(Juz.) Distr. ，29.06.1931 ，A. Lind (Holotype: LE). Shimizu in J. Fac. Tex t. Sci. Techno l.， Shinshu Univ. Distribution: steppe zone from Altai mts. in the 10 ，Se r. A ， 26: 21 (1 961). East to Austria and Hungary in the Wes t. F. F. ulmaria (L.) Maxim. subsp. megalocarpa (J uz.) Habita t: meadows and meadow steppes. Schanzer Schanzer in Bull. Soc. Na t. Moscou 94: 63 (1 989). Selected examined. specimens Moravia merid. ， ab opp. Lednice ，Vichurek (E); Voronezh gub. ，Pavlovsk ，12.01.1920 ， F. F. ulmaria auct. non Maxim. quoad pl. ex Smirnova Smirnova (LE); Volgograd Prov. ，Don valley near Kr emenskaya ， Transcaucasia ， Turcia et Ir an: Sch 凸nb.- Tem. in Rech. 25.06.1991 ，Klinkova & Schanzer (MHA); Crimea ，Chatyrdag ， 1-3.08.1894 ， Alexeenko (LE); Moscow gub. ，Tzaritzyno ， f. ， F l. Iran. 66: 122 (1 969) - Pesmen et Chamberlain 25.08.1897 ，Grigoriev (LE); S. Urals ，Cheliabinsk prov. ，Karagai in in P. Davis ， Fl. Tur k. 4: 29 (1972). forest ，3.06.1955 ，Igoshina (LE) ，Kazakhstan ， near Boluan ， 25.07.1956 ，Leontieva 245 (LE); ad fl. Irt ysh circa urb. Omsk ， Spiraea Spiraea ulmaria auc t. non L.: Boiss. ， Fl. O r. 2: 69 30.06.1885 ，Golde (LE); Altai Pr ov. ， Kamensky Distr. ，Barsuchie (1872). (1872). lake ，6.08.1954 ，Alexandrova & Guricheva (LE). Type: Asia Minor ，Iyldisdag ，Wiedemann This species is well distinguished from F. ulmaria

(Holotype: (Holotype: LE). by its cobwebby stems ，more sharply se 町 ate ， usually Distribution: Distribution: Transcaucasia ，Turkey ，NW Iran. snow-white tomentose beneath and much smaller leaf Habitat: Habitat: banks of mountain streams. segments ，a rosette usually remaining at the genera- Selected Selected specimens examined. Transcaucasia ，Armenia ， tive stalk and numerous (9-14 instead of 5-8 in F. dist r. Nor-B 吋Azet ，日. Adiaman-tshai ，26.08.1928 ，Zedelm 吋er ulmari α) ， sinuate hairy in upper p紅白 of ventral su- & Geideman (LE); Daralagoz ，Teke- Dondurak mts. ，30.07.1931 ， Karjagin Karjagin & Safiev (LE); Alagoz ，Kosha-bulag ，31.07.1932 ，E. tures achenes ， attached to a much more developed & N. Busch (LE); Caucasus Minor ，Borzhomi dist r.， Bakuriani ， than than in F. ulmaria c訂 pophore. They also differ eco- 22.07.1961 ，Ivanina & Wolk (LE). Persia ，Prov. Kordestan ，

Kuh-e 四 Chenel Chameh ， 44 km from Marivan ，7.07.1971 ， logically: e.g. ， in the Lower Don River Region in Lamond Lamond 4596 (E). Turkey ，Ovacik ，A7 ，29.07 .1 957 ，Davis & central central European Russia F. stepposa occupies mead- Hedge Hedge 31438 (E); Tatvan ，B9 ，26.07 .1 966 ， Tong 52 (E); Kars ， Sarikamis ，23.08.1957 ，Davis & Hedge 32642 (E); B8 ，Erzurum ， ows ， while F. ulmaria is restricted to alder swamps. Palandoken Palandoken Da. ，27.07.1966 ，Davis 47367 (E). Yet they sometimes hybridize in intermediate habi- This This taxon resembles F. ulmaria very much and tats ， such as forest glades ， at the northern limit of F. has has been neglected for a long time. Yet it is separable stepposa stepposa distribution ，and are usually confused. from the latter rather well both at flowering and 14. 14. (L.) Maxim. in Acta Horti fruiting fruiting stages and is quite distinct geographically. Petropo 1. 6: 251 (1 879) - Juz. ， F 1. URSS 10: 284 13. 13. Filipendula stepposa Juz. ， F l. URSS 10: 617 (1941) (1941) - Shimizu in J. Fac. Tex t. Sci. Techno l.， (1 941). Shinshu Shinshu Univ. 10 ，Se r. A ， 26: 21 - P. Ball in Tutin et F. F. ulmari α( L.) Maxim. subsp. pickbaueri (Podp.) al.， Fl. Europ. 2: 6 (1 968) ，exc l. subsp. pickbaueri Sm 吋kal in Preslia 38: 252 (1 966) - P. Ball in Tutin ， (Podp.) Smejkal - Li in Yu et al.， Fl. Reip. Pop. Sin. l. Fl. Europ. 2: 6 (1968). 37: 10 (1 985). F. F. ulmari α( L.) Maxim. va r. pickb α ueri Podp. in Filipendula ulmaria (L.) Maxim.α . tomentosa Pub l. Fac. Sci. Univ. Brno 12: 26 (1 922). (Cambess.) Maxim. op. ci t.: 252 (1 879). F. F. ulmari α( L.) Maxim. v紅 . stepposa Huber ex F. ulmaria (L.) Maxim. s. denudat α(Hayne) Janchen ，Ca t. Fl. Austr. 2: 35 (1 964). Maxim. 1. c. (1 879). F. F. ulmaria auc t. mul t.， p. F. ulmaria (L.) Maxim. var. subdenudata Fritsch Spiraea Spiraea ulmaria auc t. mul t.， p. in Abh. Zoo l. -Bo t. Ges. Wien 39: 591 (1 889). October October 1994 JoumaI of Japanese Botany Vo l. 69 No. 5 315

F. F. ulmaria (L.) Maxim. subsp. nivea Hayek ， F l. Kharkov gub. ，near Bobrovo ，5.07.1922 ，Igler (MW); Volgograd Prov. ，MedveditzaRiv. ，19.09.1974 ，Gogina& Matzenko 1446 r. St r. Exz. 8，n 356 (1 906). (MHA); (MHA); Bashkiria ，BolshoiKizilRiv. ，27.08.1955 ，Sergievskaya F. F. ulmaria (L.) Maxim. subsp. denudat α( J. et C. (LE); Tuva ，Kaa-HemDistr. ，SazimRiver. ，23.07.1978 ，Shaulo & Abakumova 1210 (MW); Krasnoyarsk Prov. ，W Sayan Mts. ， Presl) Presl) Hayek ，F 1. Steierm. 1: 872 (1909) - P. Ball in Kurtushibinsky Kurtushibinsky range ， Tikhaya Riv. ，28.07.1979 ， Shaulo & Tutin ，1. c. (1968). Saya (MW); Yakutia ，left bank of Lena Riv. ，22.07.1952 ， Kuvaev Kuvaev 13 (MW); Kr asnoyarskProv. ，StarayaIgarka ，1. 08.1947 ， F. F. denudata (J. et C. Presl) Fritsch Abh. Zoo 1. -Bo t. Tyrzhikov(MW);NMongolia ，lowerOrkhonRiv. ，23.08.1931 ， Ges. Ges. Wien 39: 591 (1 889) - Juz. ， op. ci t.， 281 (1 941). Desiatkin (LE); Evenkia ， Tunguss-Chuna Distr. ， Tunguss metheorite metheorite faU ，5.07.1961 ， Nekrasov (MHA); Lake Baikal ， Spiraea Spiraea ulmaria L.， Sp. Pl. 490 (1753). Mysovsk ，3 1. 07.1915 ，Zinserling (LE); Sikiang ，4.07.1959 ， S. S. palustris (Moench) Salisb. ，Prod r. 364 (1796). 10874 (PE). S. S. quinqueloba Baumg. ，Enum. 2: 47 (1 816). Filipendula ulmaria is rather variable ，especially S. S. denudata J. et C. Presl ， F l. Chech. 101 (1819). as to pubescence ofleaves. The most contrast variants

S. S. glauc αSchulz ，F 1. Starg. supp l. 26 (1819). with tomentose and nontomentose leaves were de-

S. S. odorata S. F. Gray ，Na t. Bri t. Pl. 2: 589 (1 821). scribed repeatingly as different varieties ， subspecies

Ulmaria Ulmaria Spiraea-Ulmaria Hill ，Hor t. Kew. 214 or even species under epithets ‘ulmaria' ，‘glauc α' ， (1 768). ‘discolor' ，‘tomentosa' ，'nivea' (tomentoseform) and U. U. pentape ω la Gilib. ， F l. Lithuan. 5: 236 (1 782). ‘denudata' (a form with leaves pubescent only along U. U. palustris Moench ，Meth. P 1. 663 (1794). veins). Intermediate forms also sometimes are recog-

U. U. ulmaria Bamh. ，Bull. To 町 ey Bo t. Club 21: 491 nized taxonomically ('subdenudat α'). As has been (1894). (1894). shown by Yapp (1912) ， these forms are stable as to

Thecanisia Thecanisia discolor (Koch) Raf. ，New F 1. Ame r. environmental conditions. Moreva (1961) cultivated 2: 2: 39 (18370. F. ulmaria at Komarov Botanical Institute Biological T. T. ulm α ria (L.) Raf. ， Sylv. Tellur. 152 (1 838). Station. She found out that the progeny of a tomentose Type: Europae pratis uliginosis umbrosis ，n 21 plant consists of both tomentose (25%) and (Holotype: (Holotype: LINN ， v. photo). nontomentose (75%) forms. However ， according to Distribution: Distribution: from the Atlantic Europe to the E as Moreva (1 961) ， the pubescence of leaves is much far far as Lake Baikal and the LenaRive r. Accidentally in more variable due to variation of the environmen t. the the Russian Far Eas t. In the Atlantic North America The result of my own study of the leaves pubes- occurs occurs as an escape from cultivation and naturalized. cence variability is that there are several types of Habitat: Habitat: floodplains of small rivers and creeks ，wet nontomentose leaf pubescence ，e.g. ， with straight to meadows. Often forms monodominant stands in vast curly hairs ， sparse to dense ， along all or main nerves areas. areas. only. All of them are deliberately united under the

Selected Selected specimens examined. British Isles ，Buckingham- epithet ‘denudat ゲ.， There is a smooth transition be- shire ，Slapton ，12.09.1977 ， Jury (お 1HA). Zealand ， near tween them and the tomentose form. Three major Haraldsted ，3.08.1971 ，Svedensen (MHA). Norvegia ，Doverfjell r.， pr.， Kongsvoll ，6.09 .1 973 ，Skvortsov (MHA). Suecia ，delta fl. groups of populations with different frequency of the Umea ，27.09 .1 973 ，Skvortsov (MHA). Fennia ， par Sysma ， tomentose form exist over the F. ulmaria range 06.1960 ，Lehtola (MHA). Switzerland ，Evoline ，20.07.1959 ， Comte Comte 3199 (MHA). Bohemia ， ad riv. Botic ， haud procul (Schanzer 1989b): European (the Atlantic Coast to Praham ，10.07.1958 ， Nitka (民 1HA). Romania ，Crisana ， the the Urals) with high frequency of ‘ denudata" form; 2.08 .1 937 ，A. & V. Borza(MHA). Smolenskgub. ，BelskDistr. ， near near Meshkovo ，14.07 .1 926 ，Savicz (LE). Estonia ， 12-14 km Siberian (E of the Urals) with extremely low fre- SE of Tartu ，7-8.08.1986 ，Skvortsov & a l. (MHA). Russia ， quency of ‘ denudata" form; North Caucasian (dis- Kursk Kursk Prov. ，Streletzky Distr. ，10.07.1947 ，Vazinger (MHA); Lvov Lvov Prov. ， near Lvov ，22.06.1946 ， Dvorakovsky (MW); junct from the main range) wi 出 rather high frequency 316 316 植物研究雑誌第69 巻第5 号 平成 6 年 10 月 of of unusuallong pilose pubescence along leaf veins. Logstor ，15.07.1969 ，Jeppesen & Pedersen (LE). England ， Newark ，23.06.1890 ， Fisher (LE). Venetia ，20.06.1904 ， G. Ser. Filipendulae. Pampanini (LE). Fl. Suecica ，Smaland ，20.06.1918 ， Kohler P 紅白 of perianth (5)6(7); achenes sessile ，laterally (LE). Polonia ，prov. Posnaniensis ，N 紘ro ，7.08.1923 ，Wisniewen (LE). (LE). Turkey ，Istanbul ，Alemdag ，29.05.1962 ，Demiriz 4942 attached attached to well developed c紅 pophore ， appressedly (E); Kars ，Sarikamis to Karakurt ，15.07.1966 ，Davis 46526 (E). pubescent pubescent throughout their surface ， densely along Ir an ，near Aliabad ，14.07.1971 ，Lamond 4844 (E). Ukraine ， E. C 紅 pathians ，Mukachevo ，27.06.1946 ， Popov (LE). Lithuania ， sutures ， not twisted; leaves multifidous ，never Kaunasdis 江， Kampishkes ，21.06.1956 ，Pipinis (LE). Belorussia ， tomentose. tomentose. Mozyr distr. ，Pripiat Riv. ，24.07.1928 ，Priakhin (LE). Middle Urals ，Sverdlovsk prov. ，Sysert Distr. ，10.07 .1 939 ，Sokolova 15. 15. Filipendula vulgaris Moench ，Meth. Pl. 663 (LE); Orenburg Prov. ，Il ek distr. ， N. of Viazovyi ，2.07.1928 ， (1794) (1794) -Hy l. in Uppsala Univ. Arsskr. 7: 196 (1 945) Borisova 374a (LE); Tobolsk gub. ， Tiumen distr. ，Li pchinskoe ， 28.06.1916 ， Gorodkov (LE); ad fl. Irt ysh circa urb. Omsk ， - P. Ball in Tutin et al.， F l. Europ. 2: 6 (1968) - 3.06.1884 ，Golde(LE);Altai ，Zmeinogorsk ，18.05.1901 ，Kr ylov Sc 凶 nb.-Tem. in Rech. f. ， F l. Ir an. 66: 122 (1 969)- (LE); Biysk ， Katun Riv. ，12.07.1903 ，Pokrovskaya (LE). Pesmen et Chamberlain in Davis ， F l. Turk. 4: 29 Filipendula vulgaris varies ov 町 its range mainly (1972). (1972). as to its overall dimensions and density of leaf pubes- F. F. hexapetala Gilib. ， F l. Lithuan. 2: 237 (1 781) ， cence ， but no clear geographic pattem can be distin- nom. i1l eg. - Maxim. in Acta Horti Petropo l. 6: 247 guished. (1 879) - Juz. ， F l. URSS 10: 286 (1941) - Shimizu in J. J. Fac. Tex t. Sci. Techno l. Shinshu Univ. 10 ， Ser. A ， Natural interspecific hybrids 26: 26: 22 (1 961). 1. F. x purpurea Maxim. in Acta Horti Petropo l. 6: F.filipendula F.filipendula (L.) Voss. in Vilmorin ，Blumengart. ， 248 (1879) - Shimizu in J. Fac. Tex t. Sci. Techno l. ed. ed. 3， 1: 240 (1896) - Rydb. ，North Ame r.日 .22:267 Shinshu Univ. 10 ， Ser. A ， 26: 10 (1961) ー Ohwi ， F l. (1 908). Jap. 537 (1 965). Spiraeafilipendula Spiraeafilipendula L.， Sp. Pl. 490 (1753). Spiraeap α lmata auc t. non Pall.: Thunb. ， F l. Jap. S. S. tuberosa Salisb. ，Prod r. 364 (1796). 212 (1784) ， p.

S. S. vulgaris (乱10ench) S. Gray ， Nat. Arr. Brit. P l. 2: Type: in Japonia ， ubi tamen semper cultam tantum 588 (1 821). vidi ， ita e Nagasaki ，Simoda ，Hakone ，Jokohama ， Ulmaria Ulmaria filipendula (L.) H i1l， Hort. Kew. 214 Hakodate. (Holotype: LE). (1 768). Exactly ， the ‘holotype' consists offour specimens Type: in pratis prope Frankenberg vor dem mounted on a single shee t: 1. a leaf of F. camtschatica Burgwald. The location of the type specimen is un- without any label; 2. n. 69 in montibus Hakone ，prope known. The type specimen of Spiraea filipendula is Foudgi- Yama ，Yaponia ，coll. Yanaka et Y okochima kept kept in LINN (Holotype ， v. photo). exeunto vere 1864 ，Maximowicz ， Iter Secundum ，

Distribution: Distribution: from the Atlanti C' coast of Europe Spiraea palmata Thunb.' (true F. X purpurea); 3. and the Atlas mts. in N. Africa eastward to the Altai 'Spiraeapalmata Th unb. ，fl. purpureis ，Simoda 1855 ， mts. mts. Accidentally in Primorsky Prov. of the Russian Japan ， capitain Jolkin' (true F. X purpurea); 4. Far East and in the Atlantic North America. 'B ipedalis ，fl. rufopurpurei ，Japonia ，Hakodate ，Ikabi ， Habita t: meadows and steppe ， subalpine mead- culta in hortis rerticanis ，12-24.10 .1 861' (stipules ows. ows. auriculate as in F. X auriculat α). These are the only Selected Selected specimens examined. Maroc ，Djebel Azidhza ， known Maximowicz's specimens of F. X purpure α prov. prov. de Demnat ， Jbrahim (LE); Bori de Boulogne ，May ， and 1 believe they should be regarded as a holotype of Parseval ・Grandmaison (LE); Grands-Malades pris de Namur ， 1867 ，Thielens & Devos (LE). Fl. Jutlandica exs. 537 ，NE of this name. October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 317

Distribution: Distribution: central and southem Japan ， in culti- Selected specimens examined. China ，Heiloungkiang ，up- per per Amur ，near Malungtiang ，1 1. 07.1950 ， Chu-Yu-Chang & vation vation and escaped. Chao-Ta-Chang 101 (LE). Russia ，Bl agoweshchensk (middle Selected Selected specimens examined. Japan ，Oizumi ，Nerima-ku ， Amur) ，2.07.1891 ，Korzhinsky (LE); Amur Prov. ，Ar khara Tokyo Tokyo Pref. ， Makino (MAK); ins. Jesso ，circaHakodate ，Albrecht Distr. ，11.06.1959 ， Shumova (MHA). 1861 1861 (LE); Pref. Nagano ，around Shi nk:吋 iTemple ，2.08 .1 973 ， These hybrids are rather variable in leaf shape Shimizu Shimizu (SHIN); Pr ef. Niigata ，Tainaigawa ，M t. Iide ，22.07.1965 ， Togashi Togashi & Yamazaki (GH). betweenF.palmata andF.α ngustilob αand are easily This This taxon is the most probably a horticultural distinguished by the unstable mosaic combinations of hybrid. hybrid. F. camtschatica ， F. multijuga and F. parentalleaf ，stem and achenes characters.

glaberrima 訂 e its putative parents. 4. F. palmata (Pal l.) Maxim. x F. camtschatica 2. 2. F. x auriculata (Ohwi) Kitam. et Murata in Acta (Pall.) Maxim. Phytotax. Phytotax. Geobo t. 20: 199 (1 962) -Ohwi ，F l. Jap. 527 Distribution: sometimes occur in southern (1965). (1965). Kamtschatka and central Sakhalin. F.purpur ωMaxim. var. auriculata Ohwi ，Fl. Jap. Specimens examined. Russia ，Kamchatka ，Kronotzky re- serve ， Bogachevka Riv. ，10.08.1982 ， Kuvaev 333-9 (MW); 647 647 (1 953) - Shimizu in J. Fac. Text. Sci. Techno l. Milkovo ， by Kamchatka Riv. ，14.07.1967 ， Mazurenko & al. Shinshu Shinshu Univ. 10 ， Ser. A ， 26: 10 (1 961). (MHA); Elizovo ，12.08.1960 ， Egorova & Woroschilow 9767 (MHA); (MHA); Sakhalin ，N ofRoshchino ，17.07.1971 ，Egoro v' a 5324 F. F. camtschatica (Pal l.) Maxim. x F. multijuga (MHA). Maxim. Hybrids are intermediate in morphology between Type: Matsunoyama-mura ， Pref. Niigata ， T. thep 紅 ental species. Haphazardly they occur in mixed Aizawa ， 06.1950 (Holotype: TNS ， n. v.). populations. Distribution: Distribution: Japan ，middle Honshu. 5. F. palmata (Pall.) Maxim. x F. ulmaria (L.) Selected Selected specimens examined. Japan ，Pref. Niigata ， Maxim. Tokimizu ，10.06.1976 ， Shimizu & Kinoshita (SHIN); Mino Pr ov. ，20.07.1922 ，Shiota (GH); Toyama Pref. ，Asahi-machi ， Distribution: very rarely occur in central Siberia. 30.07 .1 958 ， Kanai (GH). Mentioned by Popov (1 957). 1 have seen two

Filipendula Filipendula xα uriculat αis， intermediate between specimens that resemble F. ulmaria in vegetative F.c αmtschatica andF. multijuga ，andratherunstable morphology slightly deviating towards F. palmata in in in its characters ，and is distributed in the 訂 ea ofthe leaf shape ， but have straight ciliate achenes: ‘Tom ， overlap overlap of their ranges. Kondoma 2 1. Herb. Fischer (LE); Ust-Kamo ，Evenk 3. 3. F. x intermedia (Glehn) Juz. ， F l. URSS 10: 284 National Distr. ，Krasnoyarsk Prov. ，Podkamennaya (1 941) - Li in Yu et al.， F l. Reip. Pop. Sin. 37: 9 Tunguska Riv. valley ，meadows ， 16.07.1938 ， (1985). (1985). Vershikin (MW)'. Spiraea Spiraea digitata Willd. v訂 . intermedia Glehn in 6. F. ulmaria (L.) Maxim. x F. stepposa Juz. Acta Horti Petropo l. 4: 37. Distribution: occur haphazardly along the N limit F. F. angustilob α(Turcz.) Maxim. x F. palmata of the F. stepposa range ，e.g. ， in the S. Urals and the (Pall.) (Pall.) Maxim. Middle V olga Region. Type: Type: prope Kyran in Dahurie 1830 (Holotype and isotypes: isotypes: LE). 1 am grateful to Prof. A. K. Skvortsov (Main Distribution: Distribution: NEMongolia ，Ar gun andAmurRiver Botanical Garden ， Russian Academy of Sciences ， valleys. valleys. Moscow) his for guidance and encouragement during Habita t: wetfloodplainmeadows ， inhybrid swarms the course of this work ，which is actually a part of my together together with the parental species. Candidate Science (Ph. D.) thesis. 1 also wish to thank 318 318 植物研究雑誌第69 巻第5 号 平成 6 年 10 月 the the curators of all above listed Herbaria for the possi- Vostochnogo Sayana (A karyological study of the East Sayan flora species). pp. 19 --4 8. Flora Pribaikalia. Nauka ， bility bility of visiting or borrowing the materials ， Dr. Peter Novosibirsk (in Russian). Boyce (Royal Botanic Gardens ，Kew) and Dr. Jin Laane M. M. 1969. Further chromosome studies in Norvegian vascular vascular plants. Blyttia 27: 5-17. Murata (Botanical Gardens ，Nikko ， University of Lempiainen T. 1982. Mo 中hological and chemical variation Tokyo) for critical reading of the manuscrip t. among Eurasian populations of Filipendula vulgaris (Rosaceae). (Rosaceae). Ann. Bo t. Fen. 19: 127-146. Love A. and Love D. 1956. Cytotaxonomical conspectus of the References Ic elandic flora. Acta Hort. Gotob. 20: 65-29 1. Baker H. G. and Baker 1. 1967. Th e cytotaxonomy of Filipendul α 一一一一一 and E. Kjellqvist 1974. Cytotaxonomy of Spanish (Rosaceae) (Rosaceae) and its implications. Amer. J. Bo t. 54: 1027- plants. plants. U 1. Dicotyledons: Salicaceae- Rosaceae. Lagaskalia 1034. 1034. 4: 4: 3-32. Dorofeev P. 1. 1965. New data on Pliocene flora ofBashkiria. Maximowicz C. 1. 1879. Adnotationes de Spiraeaceis. Acta Stratigraphia Stratigraphia chetvertichny kh (antropogenovykh) otlozheniy Horti Horti Petropo l. 6: 105-26 1. Urala (The stratigraphy of the Quartemary (Antropogenous) Moreva T. A. 196 1. Some mo 中 hological and biological pecu- deposits deposits of the Urals). 191-219. Nedra ，Moskva (in Rus- liarities liarities of Fil 伊endula species cultivated in the North. Acta sian). sian). Ins t. Bo t. Acad. Sc i. URSS ，Se r. 5， 7: 182-219 (in Russian). 一一一一一 1977. The Simbugino flora. Fauna i flora Simbugino Morozov V. L. and Belaia G. A. 1988. Ecologia dalnevo ・ (Fauna and flora of Simbugino). 35-85. Nauka ，Moskva (in stochnogo krupnotravia (Ecology of the Far-Eastem tall- Russian). Russian). herb communities). Nauka ，Moskva (in Russi 組). Funabiki K. 1958. Distribution and polyploidy of angiosperms. Nakai T. 1914. Pla 抵抗novae Coreanae et Japoniae ， U. Feddes 1. 11. Northemflorao fJ apan. Kr omosomo37-38: 1268-1275. Repert. Repert. 13: 267-278. Gadella Th. W. J. and Kliphuis E. 1963. Chromosome numbers OhwiJ.1954.FilipendulaespeciesnovaJaponica.ActaPhytotax. offlowering plants in the Netherlands. Acta Bo t. Nee r1 and. Geobo t. 15: 115-116. 12: 12: 195-230. 一一一一一 1965. Filipendula Mil l. Flora of Japan. p. 537. Gleason H. A. 1963. Filipendula Mil l. The New Britton and Smithsonian In stitution ，Washington. Brown illustrated flora of the Northeastem United States Petrov Petrov O. M. 1976. Th e geological history ofthe Bering Strait and ajacent Canada. 2: 298-300. New York ，Dover Publica- in in the late Tertiary. Beringia v kainozoe (The Beringia in the tions tions 1nc. Tertiary). Tertiary). Vladivostok (in Russian). Grant V. 198 1. Vidoobrazovanie u rasteniy (Plant speciation). Polya L. 1949. Chromosome numbers in some Hungarian p. p. 153. Mir ，Moskva (Russian trans 1.). plants. plants. Acta Geobot. Hung. 6: 12 4- 137. JuelH.O.1918.Bei 住匂 e zur Blutenanatomie und zur Systematik Rydberg A. 1908. Filipendula Adans. North 、'A. merican Flora der der Rosaceen. Kung l. Svensk. Vetens k. Hand l.， 58: 1-79. 22: 22: 26 6- 267. Juzepczuk S. V. 194 1. Filipendula Adans. Fl ora URSS. 10: Savile Savile D. B. O. 1968. Parasite relationships and disposition of 279-289. Pub l. Acad. Sci. URSS ，Moskva (in Russian). Filipendula. Filipendula. Brittonia 20: 23 0- 231. Kalda A. and Roos A. 1973. Eesti floora P lantago jaFilipendula Schanzer 1. A. 1989a. On life forms of Filipendula. Bul l. Glavn. Ll ukide kromosoomide arv (Chromosome numbers of bo t. sada Acad. Nauk SSSR 151: 83-86 (in Russian). Plantago Plantago and Filipendula species of the Estonian flora). 一一一一一 1989b. On geographic variability and evolution of oristilised Fl oristilised markmed 15: 312-314 (in Estonian). Filipendula Filipendula ulmari α( L.) Maxim. and related species. Bull. Kalkman C. 1988. The phylogeny of Rosaceae. Bo t. J. Linn. Soc. Soc. Na t. Moscou ，Se r. Bio l. 94: 59-69 (in Russian). Soc. Soc. 98: 37-59. 一一一一一 1989c. Taxonomy and nomenclature of some Far- KartashovaN. N. ，M a1泊 chova L. A. andKozlovaA. A. 1974. A Eastem species of Fil 炉開 dul α(Rosaceae). Bo t. Zhum. 74: study study of chromosomes in the Priobie flora representatives. 959-962 (in Russian). 1. 1. Chromosome numbers of some plant species in Tomsk Sergievskaya E. V. 1967. Monografia roda Filipendula Adans. Region. Nauchnye doklady vysshei shkoly. Bio l. nauki 4: (A monograph of the genus Filipendula Adans.). Leningrad 11 4- 119 (in Russian). State State Univ. A manuscript of the candidate of science thesis Katz N. Y. ，Katz S. V. and Kipiani M. G. 1965. Atlas i (in (in Russian). opredelitel opredelitel plodov i semian vstrechaiushchihsia v Shimizu T. 196 1. Taxonomical notes on the genus Filipendula chetvertichnykh chetvertichnykh otlozheniiakh SSSR (An iconography and Adans. (Rosaceae). J. Fac. Text. Sci. Techno l. Shinshu keys to fruits and seeds occurring in Quartemary deposits of Univ. ， Ser. A (Bio l.)， 26: 1-30. the the USSR). Nauka ，Moskva (in Russian). Skvortsov A. K. 1968. 1vy SSSR ( of the USSR). Koidzumi G. 1909a. Notes on Japanese Rosaceae. Bo t. Mag. Nauka ，Moscow (in Russian). Tokyo 23: 165-189. Sokolovskaya A. P. 1960. Geographic distribution of polyploid 一一一一一 1909b. Filipendula M iI1. Conspectus Rosacearum pl 組 t species (a study of Sakhalin flora). Vestn. Leningrad. Japonicarum. Joum. Col l. Sci. Imp. Univ. Tokyo 34: 205. Gos. Univ. ， Ser. Bio l. 21: 42-58 (in Russi 加). Koshkarova V. L. 1986. Semennye flory torfianikov Sibiri 一一ー一一ー 1963. Geographic distribution of polyploid plant spe- (Seed floras of Siberian peats). Nauka ，Novosibirsk (in cies cies (a study ofKamchatka Peninsula flora). Vestn. Lenin- Russian). Russian). grad. grad. Gos. Univ. ， Ser. Bio l. 15: 38-52 (in Russian). Krogulievich R. E. 1978. Kariologicheskiy analiz vidov flory October October 1994 Joumal of Japanese Botany Vo l. 69 No. 5 319

一一一一一 1966. Geographic distribution of polyploid plant spe- Russian). cies cies (a study of Primorsky Province flora). Vestn. Lenin- Wulff H. D. 1938. Chromosomenstudien an der schleswig- grad. grad. Gos. Univ. ，Se r. Bio l. 3: 92-106 (in Russian). holsteinischen An giosperm- Fl ora. II. Be r. Deutsch. Bo t. 一一一一一， Probatova N. S. and Rudyka E. G. 1985. Chromo- Ges. 56: 247-254. some numbers of species of Asteraceae ，Poaceae ， Rosaceae Yapp R. H. 1912. ulmaria Spiraea L. and its bearing on the in in Primorsky Province ，Kamchtka and Sakhalin. Bo t. Zhum. problem of xeromo 叩hy in marsh plants. Ann. Bo t. (Lon- 70: 70: 12ι128 (in Russian). don) 26: 815-870. Sorsa Sorsa V. 1962. Chromosomenzahlen Finnischer Kormophyten 1. 1. An n. Acad. Sci. Fenn. ， se r. A ，4， Bio l. 58: 1-14. Appendix. A Ii st of recognized species and Stepanova Stepanova K. D. ， Belaya G. A.， Gurzenkov N. N. and Kachura natural interspecific hybrids and total number of N. N. 1973. Biomorphological ， physiological and specimens examined kariological kariological peculiarities of the Kamchatka meadowswee t. 1. F. occidentalis (S. Wats.) Howell 16 Pochvy irastitelnost merzlotnykh raionov SSSR (Soils and 2. 2. F. camtschatica (Pal l.) M 以 im. 280 vegetation vegetation of the USSR permafrost areas). pp. 16 6- 168. 3. F. glaberrima Nakai 212 Magadan (in Russian). 4. F. multijuga Maxim. 65 Sterling Sterling C. 1966. Comparative mo 叩hology of the c紅 pel in the 5. F.formosa Nakai 2 Rosaceae. Rosaceae. VII I. . Ame r. J. Bo t. 53: 459-520. 6. F. kiraishiensis Hayata 6 Tabata Tabata H. 1988. On the “Hi malaya co 汀 idor". Acta Phytotax. 7. F. tsuguwoi Ohwi 。 Geobo t. 39: 13-24 (in Japanese). 8. F. palmata (Pal l.) Maxim. 304 Turesson Turesson G. 1938. Chromosome stability in Li nnean species. 9. F. angustiloba (Turcz.) Maxim. 79 Ann. Ann. Agric. Col l. Sweden 5: 405 --4 16. 10. F. rubra (Hill) Robins. 55 Velichkevich Velichkevich F. Y. 1982. Pleistocenovye flory vostochnykh 11. F. vestita (Wal l. ex Hook. fi l.) Maxim. 39 lednikovykh lednikovykh oblastey Vostochno-Evropeiskoi ravniny 12. F. megalocarp αJuz. 60 (Pleistocene (Pleistocene floras of eastem glacial 訂 eas of the East- 13. F. stepposa Juz. 178 European European plain). Nauka i Tekhnika ，Minsk (in Russian). 14. F. ulmaria (L.) Maxim. 452 Woroschilow Woroschilow V. N. 1960. New findings in the Soviet Far- 15. F. vulgaris Moench 290 Eastem flora and description of new plant species. Bull. F. x purpurea Maxim. 18 Glavn. Glavn. bot. sada Ak ad. Nauk SSSR38: 42-52 (inRussian). F. x auriculata (Ohwi) Kitam. et Murata 13 一一一一一 1982. Opredelitel rastenii Sovetskogo Dalnego F. x intermedia (Glehn) Juz. 96 Vostoka Vostoka (Keys to plant species of the Soviet Far East). F.palmat αxcα mtschatic α 10 Nauka ，Moskva (in Russian). F.p α lmata x ulmaria 2 一一一一一 199 1. Filipendula Mill. (Rosaceae) species from the F. ulmaria x stepposa 5 Soviet Soviet Far Eas t. Bul l. Soc. Na t.お 1oscou 96: 125-128 (in

シャンゼル1. A.: シモツケソウ属(バラ科)の分 類学的再検討 シモツケソウ属の分類学的再検討を行った. こ 進化を遂げてきたらしい. この属のそれぞれの種 の属は 15 種からなり，北半球の温帯域に広く分布 の分布や変異のパターンを解析し，北アメリカか する.形態的形質と分布のデータをもとにすると， ら東アジアへ，そしてヒマラヤやシベリアを経由 この属には 4 つの節が認められることが明らかに した，より西方の分布域の拡大についても論議し なった. シモツケソウ属は中新世後期から最新世 た. シモツケソウ属各種の検索表と種の概念、につ にかけての移住と隔離による地理的分化によって いては本文中に記した.