10
The genus Fritillaria L. ʢLiliaceaeʣ containing 100 species, occur in West Europe to East Asia and North AmericaʢMabberley 1997ʣ . In Japan, nine species have been recordedʢNaito 2005ʣ , of which six species are listed as endangered plants in The Red Data PlantsʢYahara 2003ʣ . Fritillaria japonica groupʢNoda 1975 ; Noda and Naruhashi 1988 ; Naruhashi et al. 1997ʣ containing F . koidzumiana Ohwi are unique in that they exhibit achiasmate meiosis in pollen mother cellsʢNoda 1968, 1975ʣ , which is not ob- served in other plants in the world. Several years ago we began to examine F . koidzumiana with the aim to reveal the life history of this re- markable plant from its reproductive strategy because it is morphologically distinct from other Fritillaria species. Furthermore it is a member of spring ephemerals, shows the unique meiosis in PMCs and its taxonomically related species are endangered. Though Bakhshi Khaniki and Perssonʢ1997ʣ reported pollinators of Fritillaria , no pollinators of Japanese Fritillaria species except for F . cam- schatcensis are known until nowʢYashima et al. 1997ʣ . On the other hand, F . koidzumiana was reported as cross pollination type in breeding system by Kawano et al.ʢ2004ʣand that is con- firmedʢNaruhashi and Takata, unpublishedʣ . Therefore, it is highly meaningful study to re- veal pollinators of the plant. Fritillaria has a capsule which is upright and dry in matured time, and flat seeds with a wing are dispersed by wind. However, plants of Japa- nese F . japonica group are not upright and not dry up in matutred capsules and have somewhat globose seeds without a wing. Supposing that morphological characters, we can speculate no wind dispersal of the plant. Erythronium japonicum belonging to the Lili- aceae, is one of the famous spring ephemerals and growing with F . koidzumiana and blooming the same time, i.e., the early to the middle April in low elevation and the late April to early May in rather higher elevation in mountains in Toyama Prefecture. Both species are perennial herbs of the temperate deciduous forests of Ja- pan. The former, E . japonicum has seeds accom- panied by small elaiosomes and is reported to be myrmecochorousʢKawano et al. 1982ʣ . Kawano et al.ʢ2004ʣsuggested that the seeds of F . koidzumiana have elaiosomes and the possibility of dispersal seeds by ants. Doubtless an append- age of the seed of Fritillaria seems to be an elaiosome. Therefore, we expected to observe the seeds of F . koidzumiana to be moved around by ants. The purpose of this paper is to report on the pollinators and dispersal insects of seeds in F . koidzumiana . Materials and methods Pollinators were observed in the peak of flow- ering at four sites, see Table 1. Visiting insects were captured by net after confirmation of in- sects climbing into flower. Pollen on the insect body are examined at laboratory after capture. As noted previously, the seeds of Fritillaria are accompanied by small elaiosomes, and thus are assumed to be susceptible to myrmecochory. In order to know the dispersibility of Fritillaria seeds by ants, seeds were presented to ants in the right neighboring place of the habitats of the Journal of Phytogeography and Taxonomy 54 : 57-63, 2006 !The Society for the Study of Phytogeography and Taxonomy 2006 Naohiro Naruhashi 1 , Yuko Takata 1 and Hisashi Negoro 2 : Pollinators and dispersing insects of seeds in Fritillaria koidzumiana ʢLiliaceaeʣ 1 Department of Biology, Faculty of Science, University of Toyama, Gofuku 3190, Toyama 930ʕ8555, Japan ; 2 Toyama Science Museum, Nishinakano-machi 1ʕ8ʕ31, Toyama 939ʕ8084, Japan ʵ 57 ʵ

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Page 1: Naohiro Naruhashi , Yuko Takata and Hisashi Negoro ...phytogeogratax.main.jp/site/wp-content/uploads/2018/02/...paper no seeds of Japanese Fritillaria species are shown, because of

The genus Fritillaria L.(Liliaceae)containing100 species, occur in West Europe to East Asiaand North America(Mabberley 1997). In Japan,nine species have been recorded(Naito 2005), ofwhich six species are listed as endangered plantsin The Red Data Plants(Yahara 2003).

Fritillaria japonica group(Noda 1975 ; Nodaand Naruhashi 1988 ; Naruhashi et al. 1997)containing F . koidzumiana Ohwi are unique inthat they exhibit achiasmate meiosis in pollenmother cells(Noda 1968, 1975), which is not ob-served in other plants in the world. Severalyears ago we began to examine F . koidzumianawith the aim to reveal the life history of this re-markable plant from its reproductive strategybecause it is morphologically distinct from otherFritillaria species. Furthermore it is a memberof spring ephemerals, shows the unique meiosisin PMCs and its taxonomically related speciesare endangered.

Though Bakhshi Khaniki and Persson(1997)reported pollinators of Fritillaria, no pollinatorsof Japanese Fritillaria species except for F . cam-schatcensis are known until now(Yashima et al.1997). On the other hand, F . koidzumiana wasreported as cross pollination type in breedingsystem by Kawano et al.(2004)and that is con-firmed(Naruhashi and Takata, unpublished).Therefore, it is highly meaningful study to re-veal pollinators of the plant.

Fritillaria has a capsule which is upright anddry in matured time, and flat seeds with a wingare dispersed by wind. However, plants of Japa-nese F . japonica group are not upright and notdry up in matutred capsules and have somewhatglobose seeds without a wing. Supposing that

morphological characters, we can speculate nowind dispersal of the plant.

Erythronium japonicum belonging to the Lili-aceae, is one of the famous spring ephemeralsand growing with F . koidzumiana and bloomingthe same time, i.e., the early to the middle Aprilin low elevation and the late April to early Mayin rather higher elevation in mountains inToyama Prefecture. Both species are perennialherbs of the temperate deciduous forests of Ja-pan. The former, E . japonicum has seeds accom-panied by small elaiosomes and is reported to bemyrmecochorous(Kawano et al. 1982). Kawanoet al.(2004)suggested that the seeds of F .koidzumiana have elaiosomes and the possibilityof dispersal seeds by ants. Doubtless an append-age of the seed of Fritillaria seems to be anelaiosome. Therefore, we expected to observe theseeds of F . koidzumiana to be moved around byants.

The purpose of this paper is to report on thepollinators and dispersal insects of seeds in F .koidzumiana.

Materials and methodsPollinators were observed in the peak of flow-

ering at four sites, see Table 1. Visiting insectswere captured by net after confirmation of in-sects climbing into flower. Pollen on the insectbody are examined at laboratory after capture.

As noted previously, the seeds of Fritillariaare accompanied by small elaiosomes, and thusare assumed to be susceptible to myrmecochory.In order to know the dispersibility of Fritillariaseeds by ants, seeds were presented to ants inthe right neighboring place of the habitats of the

Journal of Phytogeography and Taxonomy 54 : 57-63, 2006�The Society for the Study of Phytogeography and Taxonomy 2006

Naohiro Naruhashi1, Yuko Takata1 and Hisashi Negoro2 : Pollinatorsand dispersing insects of seeds in Fritillaria koidzumiana(Liliaceae)

1Department of Biology, Faculty of Science, University of Toyama, Gofuku 3190, Toyama 930―8555, Japan ;2Toyama Science Museum, Nishinakano-machi 1―8―31, Toyama 939―8084, Japan

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Fig. 1. Pollinators and ants dispersing seeds of Fritillaria koidzumiana. A and B : Andrena benefica visitingflower. C : Seeds of Fritillaria koidzumiana, with scale of 1 mm. D : Lasius japonicus carrying a seed. E :Aphaenogaster famelica carrying a seed. F : Formica japonica carrying a seed.

植物地理・分類研究 第 54巻第 1号 2006年 10月

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plant(see, 1, 2, 3 and 7 in Table 2). After carry-ing the seeds into their nests, the ants were cap-tured for identification. Captured insects in thepresent study are held in trust in the samplingroom of the Toyama Science Museum.

Results and discussionBees flying about, striding into bell-shaped

Fritillaria flower and sucking nectar and collect-ing pollen, were observed(Fig. 1 A and B). Thesebees are shown in Table 3. In the table a figureare a number of captured insects. For exampleon 24th of April we observed ten bees which in-

truded into the flower, five bees, however, werecaptured. Even four times and in all places An-drena benefica Hirashima(Hymenoptera, An-drenidae)was pollinator and moreover two bees,An. hebes Pérez in Yuzunoki and An. minutula(Kirby)in Tori-dam were observed as pollinator.These three Andrena are common species in Ja-pan(Tadauchi and Hirashima 1986).

Behavior of female Andrena bees when theyvisited the flowers of Fritillaria was observed tobe as follows. They landed on the outside of te-pal, intruded into the inside of tepal, theresucked nectar and afterward collected pollen

Sites, altitudeDateTime

WeatherTemperature

Toyama Pref., Toyama-shi, Yatsuo-machi,Yuzunoki, 130 m

Apr. 15, 200610 : 00―11 : 00

Cloudy13.4℃

Toyama Pref., Toyama-shi, Yatsuo-machi,Naka, 120 m

Apr. 18, 200610 : 30―11 : 00

Fine to slightly cloudy22℃

Toyama Pref., Toyama-shi, Yatsuo-machi,Naka, 120 m

Apr. 24, 20069 : 30―11 : 30

Cloudy16.4℃

Toyama Pref., Nanto-shi, Fukumits-machi,Tori-dam, 360 m

May 4, 200613 : 30―14 : 30

Fine23℃

No. Sites and altitude Date Weather and time Ants

1 Naka in Yatsuo-machi,Toyama-shi, 120 m

May 30, 2006 Fine,14 : 20―14 : 50

Formica hayashi, Aphaenogas-ter famelica, Lasius japonicus

2 Naka in Yatsuo-machi,Toyama-shi, 120 m

May 30, 2006 Fine,14 : 50―15 : 10

Lasius japonicus

3 Naka in Yatsuo-machi,Toyama-shi, 120 m

May 30, 2006 Fine,15 : 10―15 : 30

Formica japonica

4 Campus of Univ. of Toyama,Toyama-shi, 10 m

May 30, 2006 Fine,16 : 10―16 : 20

Lasius japonicus

5 Kurehayama, Toyama-shi,15 m

Jun. 1, 2006 Fine,13 : 00―15 : 00

Formica japonica

6 Naka in Yatsuo-machi,Toyama-shi, 120 m

Jun. 3, 2006 Fine,13 : 30―16 : 00

Lasius japonicus

7 Sitanomyo in Yatsuo-machi,Toyama-shi, 220 m

Jun. 5, 2006 Fine,17 : 00―17 : 30

Formica japonica

Table 1. Sites, dates, time, weather and temperature for investigation of pollination of Fritillaria

Table 2. Investigation sites, date and dispersal ants of Fritillaria seeds

1, 2, 3 and 7, right neighboring place of population ; 4 and 5, place out of population : 6, in population.

October 2006 J. Phytogeogr. Taxon. Vol. 54. No. 1

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with thorax, postical abdomen, and hind-legsclinging or moving on a style. They sometimesstopped on the style and sucked nectar of tepal.After moving to apex portion of style or to apicalpart of tepal, they ultimately departed from theflower. Pollen was adhered on antical thorax ofAn. benefica and An. hebes when they made astop inside of the flower, but was not adhered insmall An. minutula. Pollen grains were clung onthe stigma when bees moved on style and de-parted from the flower. We observed the Fritil-laria pollen grains on the body(thorax, abdomenand hind-legs).

However, Lasius japonicus might be not a pol-linator though the ant agitated similarly andsucked nectar in the flower inside. Because of alarger flower size and a larger amount of nectarwe expected bumble-bees(Bombus)to act as pol-linator of Fritillaria. However we could not ob-serve their visit of the flower.

On the basis of this result, the pollinator ofFritillaria in Toyama Prefecture is believed tobe Andrena species.

Andrena is the largest genus of bees in Eng-land and active in March and April(Proctor andYeo 1973). Tadauchi(1992)reported Andrena inJapan visited flower from spring to early sum-mer and oligotropic. Spring ephemeral Fritillariakoidzumiana employing Andrena species as pol-linator is throught to do so as strategy for sur-vival.

Erythronium japonicum of the same familywhich grows with the present plant and bloomsat the same time, has many kinds of pollinators,such as Lepidoptera, Coleoptera, Hemiptera,Diptera and Hymenoptera(Kawano and Nagai1982). The difference of pollinator from Fritil-laria may be explained as that of the flowermorphology, flower scent and nectar component.

The pollinator of F . camschatcensis in Mt.Hakusan located in neighboring prefecture wasreported as blowfly, Aldrichina grahami(Diptera,Cyclorrhapha)by Yashima et al.(1997). Fritil-laria camschatcensis and F . koidzumiana aregrowing in the same geographical area, Ho-kuriku District in Japan Sea side, but ecologi-cally both plants are different, because the for-mer occurs in alpine region, the latter occurs inlower mountains, lower than deciduous forest

zone. Because of severe environmental condi-tions, there are only a few insects acting as polli-nators in the alpine zone. On the other handsince F . camschatcensis is pollinated by blowfly,it must show a myophilous syndrome(somewhatevil-smelling to attract blowfly, style apex deeplyparted and greatly recurved, etc.).

Different shapes of nectar glands and differentamounts and composition of nectar in the genusFritillaria is reported well by Rix and Rast(1975)and Bakhshi Khaniki and Persson(1997).Pollinators of many species are bees or bumble-bees, those of F . crassifolia and F . graeca groupare wasps and that of F . imperalis is bird(Bakhshi Khaniki and Persson 1997).

In the case of F . koidzumiana, flower isbroadly campanulate, nodding and nectar glandsoccur in grooves running along the perianth seg-ments(Naruhashi et al. 1997 ; Rix 2005). Nonectar sugars, i.e., proportion of fructose andglucose, were reported of the present flower.Generally speaking, pollination of F . koidzumi-ana is a type of bees or bumble-bees.

The seeds of Fritillaria are nearly ellipsoidaland have an elaiosome(Fig. 1 C). The results ofthe experiment of removing seeds by ants areshown in Table 2. We observed ants hold thepart of elaiosome on a seed and carried the seedinto their nests(Fig. 1 D, E and F). The ants ob-served in natural population of Fritillaria wereLasius japonicus Santschi(Formicidae, Lasiini).Similarly, the ants observed as dispersal ant atright neighboring places of populations wereFormica hayashi Terayama and Hashimoto(For-micidae, Formicini), For. japonica Motschulsky(Formicidae , Formicini)and Aphaenogasterfamelica(Fr. Smith)(Formicidae, Plagiolepidini).

At Kurehayama out of the habitat of Fritil-laria, an experiment, how do ants carry seedswas carried out. A sample was 10 seeds collectedat Naka in Yatsuo-machi, and was left near thenest of Lasius japonicus. The time of seeds re-moval by ants was eleven minutes.

A list of ants of myrmecochores in warm-temperate zone of Japan by Nakanishi(1988)contains these four ants(described For. japonicaas For. fusca and L. japonicus as L. niger in hispaper). We consider that these four ants are fa-mous for seed dispersal. In the above mentioned

植物地理・分類研究 第 54巻第 1号 2006年 10月

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paper no seeds of Japanese Fritillaria speciesare shown, because of the plants occurring incool-temperate zone. Furthermore, Nakanishi(1988)reported on the weight of disseminule(seed)for myrmecochory plant, the lightest 0.178mg in Corydalis racemosa, the heaviest 47.537mg in Akebia trifoliata and 0.4―2 mg in the mostspecies per single seed weight. Since a seedweight of the present Fritillaria is about 5.3 mg,it seems to be heavy for carring by ant. Cer-tainly the seed of Fritillaria is considered to belarge compared with the body size of ant, espe-cially that of L. japonicus.

These four ants are common and they are dis-tributed in lowland to mountainous region fromHokkaido to Kyushu and are mainly in the edgeof forests in their habitat. Moreover their nestsare made in soil(Sonobe and Onoyama 1991 ;Terayama and Yamauchi 1991 ; Watanabe andYamane 1992). The seeds of Fritillaria are notdispersed but are prevented from being feededby other animals and from dessication after re-moval into their nests by ants.

That Fritillaria is myrmecochorous, is not re-ported in Japan(Nakanishi 1999)and even inthe world(Beattie 1983). This is the first recordof the genus on myrmecochorous plant and maybe understood as a result of a quite differenthabitat, because many species of Fritillaria inthe world occur in meadows and the present spe-cies grows in woodland. Furthermore, many spe-cies in Fritillaria in medows are dispersed bythe wind because of upright dry capsules andflat seeds with a wing(cf. Turrill and Sealy1980). On the contrary, F . koidzumiana occur-

ring in woodland is myrmecochorous because ofnodding wet capsules and wingless globose seedswith a prominent elaiosome.

We wish to extend our cordial gratitude to Mr.Mamoru Sugimoto for his help in the field workand Drs. Kiyoto Maekawa and Hiroki Nakanishifor providing literatures. Thanks are also due toDr. Madjit Hakki for his corrections of Englishon our manuscript.

ReferencesBakhshi Khaniki, G. and Persson, K. 1997. Nec-

tary morphology in south west Asian Fritil-laria(Liliaceae). Nordic J. Bot. 17 : 579―611.

Beattie, A. 1983. Distribution of ant-dispersedplants. Kubitzki, K.(ed.)Dispersal and distri-bution, pp. 249―270. Verlag Paul Parey, Ham-burg and Berlin.

Kawano, S., Hiratsuka, A. and Hayashi, K. 1982.Life history characteristic and survivorship ofErythronium japonicum. Oikos 38 : 129―149.

Kawano, S, Masuda, J. and Hayashi, K. 2004.Fritillaria koidzumiana Ohwi(Liliaceae).Kawano, S.(ed.)Life history monographs ofJapanese plants. Spring Plants. vol. 1. pp.17―24. Hokkaido University Press, Sapporo.(inJapanese with English summary)

Kawano, S. and Nagai, Y. 1982. Further obser-vations on the reproductive biology ofErythronium japonicum(L.)Decne.(Liliaceae).J. Phytogeogr. Taxon. 30 : 90―97.

Mabberley, D. J. 1997. The plant book, 2 nd ed.p. 289. Cambridge University Press, Cambri-dge.

Naito T. 2005. The endemic Fritillaria species of

Sites(date) Pollinators Others

Yuzunoki(4/15) Andrena hebes 4♀An. benefica 1♀

Naka(4/18) An. benefica 4♀

Naka(4/24) An. benefica 5♀ Lasius japonicus

Tori-dam(5/4) An. minutula 4♀An. benefica 1♀

Lasius japonicus

Table 3. Pollinators in each site of Fritillaria(see, Table 1)

October 2006 J. Phytogeogr. Taxon. Vol. 54. No. 1

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Japan. Curtis’s Bot. Mag. 22 : 189―196.Nakanishi, H. 1988. Myrmecochores in warm-

temperate zone of Japan. Jpn. J. Ecol. 38 :169―176.

Nakanishi, H. 1999. Ari-ni-yoru-shushi-sanpu(Dispersal seed by ants). Ueda, K.(ed.)Shushi-sanpu, pp. 104―117. Tsukiji-shokan, Tokyo.(inJapanese)

Naruhashi, N., Sato, T. and Noda, S 1997. Com-parative anatomy of flowers in the Fritillariajaponica complex(Liliaceae). J. Phytogeogr.Taxon. 45 : 1―12.

Noda, S. 1968. Achiasmate bivalent formation byparalled pairing in PMCs of Fritillariaamabilis. Bot. Mag. Tokyo 81 : 344―345.

Noda, S. 1975. Achiasmate meiosis in the Fritil-laria japonica group I. Different modes of bi-valent formation in the two sex mother cells.Heredity 34 : 373―380.

Noda, S. and Naruhashi, N. 1988. Kobaimorui-no-shubunka-ni-okeru-senshokutaibunka-no-yakuwari. J. Jpn. Genet. 63 : 616.(in Japa-nese)

Proctor, M. and Yeo, P. 1973. The pollination offlowers. 418 pp. Collins, London.

Rix, M. 2005. Fritillaria koidzumiana. Curtis’sBot. Mag. 22 : 185―188.

Rix, E. M. and Rast, D. 1975. Nectar sugars andsubgeneric classification in Fritillaria. Bio-chem. Syst. Ecol. 2 : 207―209.

Sonobe, R. and Onoyama, K. 1991. Formica. TheMyrmecological Society of Japan(ed.). A guidefor the identification of Japanese ants(II). pp.30―35. The Myrmecological Society of Japan,Tokyo.(in Japanese)

Tadauchi, O. 1992. Hanabachi-to-kenkashokubutsu-no-kyoshinka “Coevolution of bees and phan-erogams”. Kagaku-to-seibutsu 30 : 300―304.(in Japanese)

Tadauchi, O. and Hirashima, Y. 1986. A list ofthe superfamily Apoidea(Hymenoptera)of Ni-igata Prefecture I. Andrenidae. Trans. EssaEntomol. Soc.(63):15―30.(in Japanese)

Terayama, M. and Yamauchi, K. 1991. Lasius.The Myrmecological Society of Japan(ed.). Aguide for the identification of Japanese ants(II). pp. 27―29. The Myrmecological Society ofJapan, Tokyo.(in Japanese)

Turrill, W. B. and Sealy, J. R. 1980. Studies in

the genus Fritillaria(Liliaceae). Hooker’s Icon.Plant. 39, part I and II. 1―280 pp. Bentham-Moxon Trustees, Kew.

Watanabe, K. and Yamane, M. 1992. Aphaenogas-ter. The Myrmecological Society of Japan(ed.)A guide for the identification of Japanese ants(III). pp. 15―19. The Myrmecological Societyof Japan, Tokyo.(in Japanese)

Yahara, T. 2003. Red data plants. pp. 1―720.Yama-kei, Tokyo.(in Japanese)

Yashima, T., Kinoshita, E. and Shimizu, T. 1997.Flowering phenology and self-incompatibilityin Fritillaria camschatcensis(L.)Ker-Gawl. J.Phytogeogr. Taxon. 45 : 129―133.

(Received July, 27, 2006 ; accepted September30, 2006)

鳴橋直弘1・高田由子1・根来 尚2:ユリ科コシノコ

バイモの送粉昆虫と種子散布昆虫

世界の植物の中で,花粉作成の減数分裂時に,相同染色体がキアズマを作らない唯一の植物群であるコバイモ(Fritillaria japonica group)は,8種からなり,その内コシノコバイモ(Fritillaria koidzu-miana)以外は絶滅危惧植物に指定されている。それ故早急にこの群の生活史の解明が急がれている。コシノコバイモは産地もまた個体数も多く実験に適した植物であり,筆者らは数年前よりこの植物を生殖戦略という面から分析している。この植物が他殖であることを観察(河野他 2004;鳴橋・高田 未発表)したが,送粉昆虫は不明だったので,2006年 4月 15日から 5月 4日に 4回,富山県の 3カ所の生育地において観察・調査した。その結果,4回とも共通して膜翅目ヒメハナバチ科のウヅキヒメハナバチ(Andrena benefica)が訪花して,花粉を媒介していたことを観察した。それ以外に同属のヤヨイヒメハナバチ(An. hebes)とマメヒメハナバチ(An. minutula)も送粉昆虫であることを確認した。コシノコバイモの種子にエライオソームが観察されアリ散布が期待されたので,コシノコバイモの個体群内 1カ所での観察と生育地ではあるが個体群外 4カ所,および生育地外 2カ所で,種子をアリが運ぶかどうかの実験を行った。その結果,ハヤシクロヤマアリ(Formica hayashi),クロヤマアリ(F. japonica),ア シ ナ ガ ア リ(Aphaenogasterfamelica),トビイロケアリ(Lasius japonicus)が種子を口にくわえて運ぶ姿を観察した。本種が属するバイモ属は,欧州,アジア,北米と北半球に広く分布し,世界に約 100種存在するが,種子のアリ

植物地理・分類研究 第 54巻第 1号 2006年 10月

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散布はこの属での最初の報告である。本属の多くの種は草原性で,それらの種子は扁平で翼を持ち,風散布と考えられるのに対して,コバイモは森林性で,種子は楕円体に近く翼がなく,エライオソームを持ち,アリ散布である。本実験において捕獲した昆虫

は,富山市科学文化センターに保存されている。(1〒930―8555 富山市五福 3190 富山大学理学部生物学科;2〒939―8084 富山市西中野 1―8―31富山市科学文化センター)

October 2006 J. Phytogeogr. Taxon. Vol. 54. No. 1

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植物地理・分類研究 第 54巻第 1号 2006年 10月

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表紙図の説明:描かれた植物は,ユリ科バイモ属コシノコバイモで,山形県から石川県の日本海側とまれに岐阜県,愛知県,静岡県にも分布する。北陸では産地も個体数も多く,他地域と比べ大形である。本学会が金沢で生まれ北陸を中心に発展してきたことから,この北陸の植物を表紙に使用した。図は梅林正芳氏が描いたもの。

目 次

論文

岩松佳代・沖津 進:富士山高山域の異なる土壌粒径上に生育するオンタデの地下形態 ……………………1

花井隆晃・渡邊幹男・芹沢俊介:ミゾソバ(タデ科)の分類学的再検討 (2)滋賀県浅井町および東京都高

尾山における群構成 …………………………………………………………………………………………………7

増田 優・ファリニアイナ ルシエン・近藤勝彦:マダガスカル産ラン科 Gastrorchis と Phaius の分子系統

学的関係 ……………………………………………………………………………………………………………15

芹沢俊介:淡黄色花タンポポの分類 ………………………………………………………………………………21

中西弘樹・中西こずえ・岩城太郎:サキシマフヨウの花の形質と変異,特にフヨウとの比較において …27

鳴橋直弘・和田賢次:トクシマコバイモ Fritillaria tokushimensis(ユリ科)は自然雑種と考えられる …35

短報

岩松佳代・沖津 進:富士山高山域の異なる土壌粒径上に生育するオンタデ個体群のサイズ分布 ………45

山田直樹:シーボルト入手の植物標本帳Herbarium Medici Jedoensisにあるスミレの標本 ………………51

鳴橋直弘・高田由子・根来 尚:ユリ科コシノコバイモの送粉昆虫と種子散布昆虫 ………………………57

池田 博・津坂真智子・兼子伸吾・狩山俊悟:ミチノクフクジュソウ(キンポウゲ科)の新産地と核型 …65

ノート

梅林正芳・加藤 真・太田道人・本谷二三夫・中野真理子・木下栄一郎:マイヅルテンナンショウ(サトイモ

科),ハイハマボッス(サクラソウ科),ミズニラ(ミズニラ科)の富山県内における新産地 …………71

大原隆明・高木末吉・安嶋 隆・源内伸秀:ススヤアカバナ Epilobium parviflorum を日本のフロラに追加

する …………………………………………………………………………………………………………………74

村中孝司:鬼怒川(栃木県)河川敷に生育する絶滅危惧植物ムラサキセンブリ Swertia pseudochinensis(リ

ンドウ科) ……………………………………………………………………………………………………………78

新刊紹介 ………………………………………………………………………………………………………………81

植物関連雑誌のタイトル紹介 ………………………………………………………………………………………86

会記 ……………………………………………………………………………………………………………………92

CONTENTS

ArticlesKayo Iwamatsu and Susumu Okitsu : Root morphology of Polygonum weyrichii var. alpinum growing

on different soil particle sizes on the alpine area of Mt. Fuji ………………………………………………1Takaaki Hanai, Mikio Watanabe and Shunsuke Serizawa : Taxonomic studies of Polygonum thunbergii

s. lat.(Polygonaceae)(2)Subpopulation structures in Azai-cho(Shiga Prefecture)and Mt. Takao(Tokyo Prefecture) …………………………………………………………………………………………………7

Yu Masuda, Lucien Faliniaina and Katsuhiko Kondo : Phylogenetic analysis of Malagasy Gastrorchisand Phaius(Orchidaceae)based on internal transcribed spacer(ITS)sequence …………………15

Shunsuke Serizawa : Classification and nomenclature of the pale-yellow-flowered dandelions in Japan…………………………………………………………………………………………………………………………21

Hiroki Nakanishi, Kozue Nakanishi and Tarou Iwaki : Floral characters and their variations of Hibis-cus makinoi(Malvaceae), with special reference to comparison with H . mutabilis …………………27

Naohiro Naruhashi and Kenji Wada : Fritillaria tokushimensis(Liliaceae)is thought to be a naturalhybrid ………………………………………………………………………………………………………………35

Short communicationsKayo Iwamatsu and Susumu Okitsu : Size distributions of Polygonum weyrichii var. alpinum growing

on different soil particle sizes in the alpine area of Mt. Fuji ………………………………………………45Naoki Yamada : Four specimens of Viola included in the plant album “Herbarium Medici Jedoensis”

obtained by von Siebold in 1827 ………………………………………………………………………………51Naohiro Naruhashi, Yuko Takata and Hisashi Negoro : Pollinators and dispersing insects of seeds in

Fritillaria koidzumiana(Liliaceae) ……………………………………………………………………………57Hiroshi Ikeda, Machiko Tsusaka, Shingo Kaneko and Shungo Kariyama : A new locality of Adonis mul-

tiflora(Ranunculaceae)and its karyomorphology …………………………………………………………65

Miscellaneous notesMasayoshi Umebayashi, Makoto Kato, Michihito Ohta, Fumio Mototani, Mariko Nakano and Eiichiro

Kinoshita : New localities of Arisaema heterophyllum(Araceae), Samolus parviflorus(Primulaceae)and Isoetes japonica(Isoetaceae)in Toyama Prefecture …………………………………………………71

Takaaki Oohara, Suekichi Takagi, Takashi Ajima and Nobuhide Gen-nai ; Newly distributed Epilo-bium parviflorum in Japan ………………………………………………………………………………………74

Takashi Muranaka : Present status of the endangered plant Swertia pseudochinensis(Gentianaceae)in the floodplain of the Kinu River, Tochigi Prefecture ……………………………………………………78

Book reviews ………………………………………………………………………………………………………81

Titles of articles in other botanical journals ………………………………………………………………86

Announcements ……………………………………………………………………………………………………92

編集委員会 (Editorial Board)

編集委員長 (Editor-in-Chief) 岩坪 美兼 (Yoshikane Iwatsubo, Toyama)

主任編集委員 (Managing Editor) 和田 直也 (Naoya Wada, Toyama)

福嶋 司 (Tukasa Hukusima, Tokyo) 木下栄一郎 (Eiichiro Kinoshita, Kanazawa)

中田 政司 (Masashi Nakata, Toyama) 高橋 弘 (Hiroshi Takahashi, Gifu)

綿野 泰行 (Yasuyuki Watano, Chiba) Madjit I. Hakki (Berlin, Germany)

Frederick H. Utech (Pittsburgh, U.S.A.)

Page 10: Naohiro Naruhashi , Yuko Takata and Hisashi Negoro ...phytogeogratax.main.jp/site/wp-content/uploads/2018/02/...paper no seeds of Japanese Fritillaria species are shown, because of

2006年 10月 31日発行 編集発行人 〒920―1192 金沢市角間町 金沢大学大学院自然科学研究科植田 邦彦

発 行 所 〒920―1192 金沢市角間町 金沢大学大学院自然科学研究科植物自然史研究室内 植物地理・分類学会

印 刷 所 〒920―0855 金沢市武蔵町 7番 10号 能登印刷株式会社

� The Society for the Study of Phytogeography and Taxonomy 2006

Volume 54. No. 1October 2006

(通巻 第 159号)

ISSN 0388-6212

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