植物研究雑誌 J. J. Jpn. Bo t. 74: 74: 296-306 (1 999)

Morphological Morphological Variation and Distribution of the the Cotyledon Areoles in Papilionoideae (Leguminosae) and Their Their Systematic Utility

b Yasuhiko ENDO a and Hiroyoshi OHASHI

aNatural aNatural History Museum and Institute ，Chiba ，Aoba-cho 955-2 ，Chuo ・ku ，Chiba ，260-8682 JAPAN; bBiological bBiological Institute ，Graduate School of Scie 附， Tohoku University ，Sendai ，980-8578 JAPAN (Received (Received on March 24 ，1999)

The cotyledon areole (= CA) was observed in 108 among 147 species examined examined in the subfamily Papilionoideae of Leguminosae. These species are in- cluded cluded in 69 genera and in 22 tribes. CA's shape ，position and size were approxi- mately mately illustrated on the mature seed embryo in each species. CAs were triangular ， circular ，oval ，oblong ， or linear ， and situated at the base ，basal-middle ，middle ，api- cal-middle ， or apex of the cotyledons. The shape and position of the CAs were usu- ally ally similar in a genus and may be useful as characters for consideration of phylo- genetic genetic relationships among the genera in Papilionoideae.

Key words: Cotyledon areole (CA) ，distribution ，Leguminosae ，morphological varia- tion ，Papilionoideae

The cotyledon areole (= CA) is a struc- 1995). In the present paper ，furthermore ，we ture of cotyledons in the subfamily aim to discuss the systematic utility in other Papilionoideae Papilionoideae of Leguminosae (Endo and Papilionoid legumes. Ohashi 1998a). It exists on the midvein of cotyledons cotyledons of mature seeds and shows ver- Materials and Methods rucae rucae by projection of epidermal cells (Endo We examined 147 species belonging to 98 and Ohashi 1997 ，1998a ，1998b ， 1999). The genera and to 23 tribes. The species exam- CAs vary in size and shape in different taxa ined are listed in Table 1. (Endo and Ohashi 1998a). The position is For observation of embryo ，we took off different different also in different taxa (Endo and the seed coat after softening mature seeds of Ohashi 1998a). The shape and position were the species mentioned above by soaking considered considered to be useful as taxonomic char- them into 50% aqueous ethanol more than acters acters for higher taxa above the rank of gen- one day. Outlines of the embryos and CAs era era (Endo and Ohashi 1998a). In a previous were sketched with a dissecting microscope paper (Endo and Ohashi 1998a) ，' we dis- attaching camera lucida. cussed cussed systematic utility of the CAs to dis- tinguish tinguish Vicioid clade (sensu Sanderson and Results and Discussion Wojciechowski 1995) from tribe Presence or absence of the CA in the spe- Carmichaelieae Carmichaelieae in temperate legume clade cies examined was presented in Table 1. In (sensu Sanderson and Wojciechowski the table ，tribes were a町 anged in accordance

-296 ー October October 1999 Journal of Japanese Botany Vo l. 74 No. 5 297

Table 1. Distribution of the cotyledon areole in Papilionoideae

Tribe Tribe Species examined and voucher specimen (1) CA(2) Fig.

Sophoreae Sophoreae Sophora flavescens Aiton: South Korea: T. Nemoto et al. 96911005 (TUS) + Cladrastis Cladrastis lutea (F.Michx.) K. Koch * Maackia amurensis Rup r. ex Maxim.: South Korea ， lnchon: H. Ohashi et al. in 1996 (TUS) - Sophora tomentosa L.: Australia: H. Ohashi et al. in 1998 (TUS) Abreae Abrus precatorius L. +* 2 Affiorpheae Affiorpheae Amorpha fruticosa L. * Dα lea alopecuroides Willd. * Millettieae Millettieae Tephrosia obovata Mer r. +* 3 Tephrosia Tephrosia purpurea (L.) Pers. +* 4 Craibia Craibia zimmermannii (Harms) Dunn * Millettia Millettia grandis (E ふ1ey.) Skeels 一*

Robinieae Robinieae Petteria rα， mentacea (Sieber) C.Presl +* 5 Robinia Robinia pseudoacacia L. +* 6 Sesbania Sesbania cannabina (Retz.) Poi r. +* 7 Sesbania Sesbania punicea (Cav.) Benth.: North America: Y. Tateishi et al. 18305 (TUS) + 8 Sesbania Sesbania sp. +* 9 Sesbania Sesbania sp.: North America: H. Ohashi et al. 961013101 (TUS) + 10 Indigoferae Indigoferae Indigofera australis Willd.: Australia: H. Ohashi et al. in 1998 (TUS) + 11 Indig o.. 作ra hirsuta L.: Philippines: T. Nemoto et al. 10647 (TUS) + 12 Indigofera Indigofera linifolia (L .f.) Retz. +* E&O 98a ，日*** Indigofera Indigofera zollingeriana Miq. +* 13 Indigofera Indigofera suffruticosa M i1l. * Indig o.. 大 ra kirilowii Maxim.: South Korea ， lnchon: H. Ohashi et al. in 1996 (TUS) 'I'11121'i'IA Phaseoleae Canavalia sp.: Australia: H. Ohashi et al. in 1998 (TUS) + TZJζU叫 Centrosema Centrosema sp.: Australia: H. Ohashi et al. in 1998 (TUS) + Clitoria Clitoria sp.: Australia: H. Ohashi et al. in 1998 (TUS) + 守 Dumasia sp.: Philippines: T. Nemoto et al. 10528 (TUS) + I00 Galactia Galactia regularis (L.) BSP.: North America: H. Ohashi et al. in 1996 (TUS) + 白 Glycine Glycine max (L.) Merr. +* YAU'I 司， Glycine Glycine sp.: Australia: H. Ohashi et al. in 1998 (TUS) + h 司 Pachyrhizus Pachyrhizus erosus (L.) Urb.: Philippines: T. Nemoto et al. 10062 (TUS) + fH 司 司・ Rhynchosia Rhynchosia volubilis Lou r.: Philippines: T. Nemoto et al. 10540 (TUS) + ''u4 Apios Apios americana Medik.: North America: Y. Tateishi et al. 18327 (TUS) Cajanus Cajanus cajan (L.) Millsp.: Philippines: T. Nemoto et al. 10063 (TUS) Cajanus Cajanus reticulatus F.Muell.: Australia: H. Ohashi et al. in 1998 (TUS) Cajanus Cajanus scarabaeoides (L.) Thouars: Philippines: T. Nemoto et al. 10648 (TUS) Dunbaria villosa (Thunb.) Makino: South Korea ， lnchon: H. Ohashi et al. in 1996 (TUS) Eη thrin αcristagalli L.: Royal Botanic Gardens in Sydney ，Australia: n.v.s. Flemingia Flemingia macrophylla (Willd.) Kuntze ex Prain: Royal Botanic Gardens in Sydney ，Australia: n.v.s.

Kennedia coccinea Ven t. ーネ Macroptilium Macroptilium lathyroides (L.) Urb.: Australia: H. Ohashi et al. in 1998 (TUS) Phaseolus Phaseolus angularis W.Wight: South Korea ， M t. Chili: H. Ohashi et al. in 1996 (TUS) Strophostyles Strophostyles helvola (L.) Britton: North America: H. Ohashi et al. in 1996 (TUS) Vigna Vigna marina (Burm.) Mer r.: Australia: H. Ohashi et al. in 1998 (TUS) DeSffiodieae DeSffiodieae Codariocalyx gyroides Hassk. +* 23 Desmodium brachypodium A.Gray: Australia: H. Ohashi et al. in 1998 (TUS) + 24 Desmodium gunnii Hook.f.: Australia: H. Ohashi et al. in 1998 (TUS) + 25 Desmodium heterocarpon (L.) DC.: Philippines: T. Nemoto et al. 10374 (TUS) + 26 Desmodium heterocarpon (L.) DC.: Japan ，Kagoshima Pref.: M. Furuse 11953 (CBM) + Desmodium paniculatum (L.) DC.: Japan ，Chiba Pref.: K. Yamamoto in 1997 (CBM) + 27 Desmodium sequax Wall.: Philippines: T. Nemoto et al. 10397 (TUS) + 28 Pycnospora Pycnospora lutescens (Poi r.) Schind l. +* 29 Tadehagi Tadehagi triquetrum (L.) H.Ohashi ssp. pseudotriquetrum (DC.) H.Ohashi +* 30 Uraria Uraria lagopodioides (L.) Desv. ex DC. +* 31 Alysicarpus Alysicarpus ovalifolius (Schum.) J.Leonard * 298 植物研究雑誌第74 巻第5号 平成11 年10 月

(1) (1) (2) ..... (3) Tribe Tribe Species examined and voucher specimen CA'-' Fig.

Cα mpylotropis giraldii (Schi 凶nd 訓1.) Schind 訓1. . * Chr バisti αobcord αtα(Poirι.)， Bakh.f 一* Dendrolobi ωum ηlum ηlbella αtω um η1 (L.) Bent 出h. 申 Desmodium caudatum (Thunb.) DC.: Japan ，Chiba Pref.: H. Yamai in 1996 (CBM) Desmodium oldhamii Oliv.: Japan ，Chiba Pref.: S. Ts 吋i in 1997 (CBM) Desmodium Desmodium podocarpum DC. ssp. fallax (Schind 1.) H.Ohashi: Japan ， Chiba Pref.: T. Tanabe in 1997 (CBM) Desmodium Desmodium podocarpum DC. ssp. oxyphyllum (DC.) H.Ohashi: Japan ，Chiba Pref.: T. Kawana in 1993 (CBM) Desmodium laxum (L.) DC. : Japan ，Chiba Pref.: H. Yamai in 1988 (CBM) Desmodium laxum (L.) DC. ssp. leptopus (A.Gray ex Benth.) H.Ohashi * Kummerowia stipulacea (Maxim.) Makino * Lespedeza Lespedeza formosana (Vogel) Koehne * Phyllodium Phyllodium pulchellum (L.) Desv. 一* Psoraleae Psoraleae Psoralea bituminosa L. +* 32 Loteae montana L. +* 33 Anthyllis Anthyllis vulneraria L. ssp.α lpestris Asch. et Graebn. +* 34 Coronilla Coronilla emerus L. +* 35 Hippocrepis Hippocrepis balearica Jacq. +* 36 Hymenocarpus circinnatus (L.) Savi +* 37 Hymenocarpus nummularius (D C.) G.Don +* 38 Lotus Lotus corniculatus L. var. japonicus Regel +* 39 Ornithopus Ornithopus sativus Bro t. +* 40 Scorpiurus Scorpiurus vermiculatus L. +* 41 持 tragonolobus maritimus (L.) Roth +* 42 Tetragonolobus Tetragonolobus palaestinus Boiss. +* 43 Tetragonolobus Tetragonolobus requienii (Mauri ex Sanguin.) Daveau +* 44 Tetragonolobus Tetragonolobus purpureus Moench. * Aeschynomeneae Aeschynomeneae Aeschynomene indica L. +* 45 Nissolia Nissolia fruticosa Jacq. +* 46 Arachis Arachis hypogea L. 本 Galegeae Alhagi maurorum Medik. +* 47 Astragalus membranaceus Fisch. +* 48 Biserrula pelecinus L. +* 49 Calophaca tianschanica Boriss. +* 50 Clianthus puniceus Banks & So l. ex Lind l. +* E&O 98a ， fl Colutea paulsenii Freyn +* 51 o.ffi cinalis L. +* 52 Galega orientalis Lam. +* 53 Glycyrrhiza Glycyrrhiza pallidiflora Maxim. +* 54 Guerdenstaedtia Guerdenstaedtia stenophyll αBunge. +* 55 Halimodendron halodendron (Pall.) Voss +* 56 Lessertia perennans DC. +* 57 Oxytropis strobilacea Bunge. +* 58 R. B r. +* 59 Sw α insonia galegifolia R. Br. +* 60 Caragana arborescens Lam. * Caragana maximowicziana Kom. * Carmichaelieae Carmichaelieae Cα rmichaelia australis R. B r. +* 61 Carmichaelia exsul F ぶ1uel l. +* 62 Chordospartium Chordospartium stevensonii Cheeseman +* 63 Hedysareae austrosibiricum B.Fedtsch. +* 64 Hedysarum boreale Nutt. +* 65 Hedysarum hedysaroides (L.) Schinz et TheI I. +* 66 Hedvsarum ussuriense I. Schischk. & Kom. +* 67 Hedysarum vicioides Turcz. +* 68 Onobrychis Onobrychis montana DC. +* 69 October October 1999 Journal of Japanese Botany Vo l. 74 No. 5 299

Tribe Tribe Species examined and voucher specimen (1) CA(2) Fig. (3)

*** 司 ハ SuUa coronaria (L.) Medik. ** +++ Iυ21 勺 Fabeae Fabeae (=Vicieae) Lathyrus sylvestris L. I 吋，， 『1 Lens esculenta Moench ・ M Pisum sativum L. +* E&O 98a ， f8 日cia americana Muh l. ex Willd. var. sinensis C. R. Gunn +* 73 Vicia Vicia amurensis Oet t. +* 74 Vicia Vicia angustifolia L. +* 75 Vicia Vicia articulata Hornem. +* 76 Vicia Vicia bザ'o lia Nakai +* 77 Vicia Vicia bithynica (L.) L. +* 78 日cia dumetorum L. +* 79 1々cia faba L. +* Viciafaba Viciafaba L.: Seeds obtained from Hor t. Bo t. Univ. Debrecen ， Hungaria: n.v.s. + 80 Vicia Vicia nigricans Hook. & Arn. +* 81 Vicia Vicia pisiformis L. +* 82 Vicia Vicia sp. (from Chile) +* 83 Cicereae Cicereae Cicer arietinum L. +* E&O 97 ，自料水 Trifolieae Trifolieae Medicago arborea L. +* 84 Melilotus Melilotus altissima Thuil l. +* E&O 98a ，f4 Ononis Ononis alopecuroides L. +ネ 85 Ononis Ononis spinosa Benth.: Seeds obatain 巴d from Bo t. Gard. of the Univ. Nonnensteeg ，Netherlands: n.v ふ+ 86 +++++++一++++************ Ononis Ononis viscosa L. 87 Parochetus Parochetus communis D.Don 88 Trifolium Trifolium clypeatum L. 89 Trifolium Trifolium miege αnum Maire 90 TrigoneUa TrigoneUa foenum-graceum L. 91 Bossiaeeae Bossiaeeae Goodia lotifolia Salisb. 92 Lamprolobium fruticosum Benth. 93 Bossiaea Bossiaea brownii Benth. 0JOynynynyny456789 Mirbelieae Mirbelieae Mirbelia speciosa Sieber ex D C. Oxylobium lanceolatum (Ven t.) Druce Podalyrieae Podalyrieae Podalyria sericea R. B r.

Crotalarieae Crotalarieae Crot α， laria sp. Crotalaria Crotalaria sp.: Royal Botanic Gardens in Sydney ，Australia: n.v.s. + +一++++牢***** Thermopsideae Thermopsideae Anagyris foetid αL， Thermopsis Thermopsis lupinoides (L.) Link Genisteae Genisteae Cytisus scoparius (L.) Li nk 100 Lupinus Lupinus arcticus S.Watson 101 Lupinus Lupinus luteus L. 102 Sp α rtium junceum L. 103

(1) (1) No data of voucher specimens of the species are same as those listed in our previous paper (Endo and Ohashi 1998a ，Table 1). n. v .s. = no voucher specimen. (2) (2) Presence or. absence of cotyledon areole (+ CA present ，- CA absent). * = data from Endo and Ohashi (l 998a). **Sulla **Sulla coronaria was treated as Hedysarum coronarium in the previous paper. (3)Figure (3)Figure in the present paper- ***For example ，E&O 98a ， f3 means Fig. 3 in Endo and Ohashi (1 998a); and E&O 97 ， f5 means Fig. 5 in Endo and Ohashi (1 997). 300 植物研究雑誌第74 巻第5号 平成11 年10 月 with with the systematic sequence by Polhill Millettieae (Figs. 3 ，4); CAs were situated (1 994). In the tribes ， species were di vided at apical-middle part of cotyledons ，and into into two groups; CA present and CA absen t. CAs' outlines were ova l. Species Species in each group were arranged by al- Loteae (including Coronilleae) (Figs. 33- phabetical phabetical orde r. The CA was observed in 44); CAs were situated at basal (Figs. 33 ， 108 108 species which are included in 22 tribes 34 ，39 ，42-44) ，basal-middle (Figs. 35-38 ， (Table (Table 1). These tribes are about 739 もof all 41) or middle (Fig. 40) part of cotyledons ， the the known Papilionoid tribes (30 tribes). and the shape was oval (Figs. 33-39 ， 41-44) Shape of CAs was illustrated in Figures or oblong (Fig. 40). The position observed 1-103. 1-103. The order of these figures follows shows no uniformity but closely to each that that in Table 1. The shape is circular ，oval ， other. And the shapes could not be divided oblong ，linear ， or triangular. Position of CAs into any types because of presence of inter- is is different in different taxa but CAs always mediate shapes. appear appear on midvein of the cotyledon blades. Fabeae (Figs. 71-83); CAs were situated To describe the position of CA we divide at the base ， and were ova l. cotyledon cotyledon into five equal parts from the base Trifolieae (Figs. 84-91); CAs were situ-

to to apex ，i.e. ，basal ，basal-middle ，middle ， 戸 ated at basal (Figs. 85 ，86 ， 88-90) or basal- apical-middle ，and apical part respectively. middle (Figs. 84 ，87 ， 91) part of cotyledons ， The shape and position of CAs are usu- and were oval (Figs. 85-90) or oblong (Figs. ally ally similar to each other among different 84 ， 91). species species included in one genus. However ， Bossiaeeae (Figs. 92 ，93); CAs were situ- there there are some variations in lndigofera and ated at basal ， and were triangular (Fig. 92) Desmodium. or oval (Fig. 93). CAs of five lndigofera species were situ- However ，some obvious differences were ated ated at apical- middle part of the coty ledons found in Phaseoleae (Figs. 14-22) ， Galegeae (Figs. (Figs. 11-13; Fig. 3 in Endo and Ohashi (Figs. 47-60) ，and Genisteae (Figs. 100- 1998a). 1998a). Approximate shapes of the CAs 103). were oval (Fig. 12; Fig. 3 in Endo and Abreae (Fig. 2) ，a part of Millettieae Ohashi Ohashi 1998a) ， oblong (Fig. 13) ， or linear (Tephrosi α(Figs. 3， 4) and Derris) ， (Fig. (Fig. 11). In the genus Desmodium ， five of lndigoferae (Figs. 11-13) ，Phaseoleae (Figs. nine nine species examined have CA (Figs. 24- 14-22) ，Desmodieae (Figs. 23-31) and 28). 28). The CAs were situated at middle part Psoraleae (Fig. 32) were considered to com- (Figs. (Figs. 25 ，28) ， at apical-middle part (Figs. pose a monophyletic group ， old world tropi- 24 ，27) ， or at adaxial edge of apical-middle cal tribes ， from molecular data (Doyle et al. part part of the cotyledon blade (Fig. 26). We 1997). Shapes of CAs were usually oval in need need further investigation to clarify varia- the group but were linear in lndigofera aus- tions tions of the characters of the CA in these tralis (Fig. 11) and Dumasia sp. (Fig. 17). two genera for consideration about intra- Position was basal (Figs. 2，32) ，middle generic generic systematic utility of the characters. (Figs. 15-17 ，19-21 ，23 ，25 ，28 ，30) ，apical- In In the following tribes ，we could not find middle (Figs. 3，4 ，11-13 ，24 ，26 ，27 ，29 ， any any obvious differences in shape and posi- 31) ， or apical part (Figs. 14 ，18 ， 22) of coty- tion tion of CAs among different genera belong- ledons. The apical position of CA is unique ing ing to the same tribe and we thought that to Papilionoid cotyledons. Such position was these these characters. might be difficult to use to observed in Galactia ，Canavali αand consider consider phylogenetic relationships among Rhynchosia of the tribe Phas t1 01eae. A part the the genera included in the same tribe: of Diocleinae in which Galactia and October October 1999 Journal of Japanese Botany Vo l. 74 No. 5 301

lll s，

16 16

(il¥ij/ れ¥ 。

Figs. Figs. 1-32. Embryos and cotyledon areoles (black-painted) of Sophoreae (1)， Abreae (2) ， M i1l ettieae (3 ，4) ，Robinieae (5-10) ， Indigoferae (1 1-13) ，Phaseoleae (1 4-22) ， Desmodieae (23-3 1)， and Psoraleae (32). Species names of the figures are listed in Table 1. 1. Scale bars = 1 mm. 302 302 植物研究雑誌第74 巻第5号 平成 11 年10 月 刊、"

58 58

Figs. Figs. 33-62. Embryos (33-55 ，57-62) ，a cotyledon (56) ， and cotyledon areoles (black- painted) painted) of Loteae (33-44) ，Aeschynomeneae (45 ，46) ，Galegeae (47-60) ，and Carmichaelieae Carmichaelieae (61 ，62). Species names of the figures are listed in Table 1. Scale bars = 1 mm. October October 1999 Journal of Japanese Botany Vo l. 74 No. 5 303

lili--40

40

11111EOO

司 I

Figs. Figs. 63-9 1. Embryos (63-83 ，86-91) ， cotyledons (84 ， 85) and cotyledon (black- areoles painted) painted) of Carmichaelieae (63) ，Hedysareae (64-70) ，Fabeae (71-83) ，and Trifolieae (84-9 1). Species names of the figures are listed in Table 1. Scale bars = 1 mm. 304 植物研究雑誌第74 巻第5号 平成11 年 10 月

li--9III-

ヲ a

Figs. Figs. 92 ー103. Embryos and cotyledon areoles (black-painted) of Bossiaeeae (92 ，93) ， Mirbelieae Mirbelieae (94 ，95) ，Podalyrieae (96) ，Crotalarieae (97 ，98) ， Thermopsideae (99) ， and Genisteae Genisteae (l 00-1 03). Species names of the figures are listed in Table 1. Scale bars = 1 mm.

Canav αlia are in c1 uded was considered to be group ，Vicioid clade (Sanderson and the the sister group of C 吋aniinae ，w hich in- Wojciechowskii 1995 ， 1996). Position and c1 uded Rhynchosia ，from chloroplast DNA shape of CAs of Fabeae and Trifolieae were data data (Doyle and Doyle 1993 ， Bruneau et al. mentioned above. CAs of the genus Galega 1994). 1994). From these facts ， the apical position were situated at basal-middle part of coty- of of CA may indicate close phylogenetic re- ledons and shape of CAs was ova l. CA of lationships lationships among these subtribes. In the old Cicereae was situated at basal part of a coty- world world tropical tribes ， the positions of CAs ledon ， and the shape was ova l. Therefore ， may be useful to consider inter-generic phy- CAs of Vicioid c1 ade were closely situated logenetic logenetic relationships. each other and the shapes could not be di- Galegeae was considered to be a vided into any types because of presence of paraphyletic paraphyletic group from molecular data intermediate shapes. We thought that it (Sanderson (Sanderson and Wojciechowski 1996 ， Doyle might be difficult to use the position and et et al. 1997). On the other hand ，genus shape of CAs to consider intra- and inter- Galega (Figs. 52 ， 53) of the tribe Galegeae ， generic phylogenetic relationships in the Fabeae Fabeae (Figs. 71-83) ，Trifolieae (Figs. 84- Vicioid c1 ade. 91) ，and Cicereae (Fig. 5 in Endo and Genus Alhagi (Fig. 47) of the tribe Ohashi in 1997) inferred a monophyletic Galegeae ， and genera Hedysarum (Figs. 64- October October 1999 Journal of Japanese Botany Vo l. 74 No. 5 305

68) ， Onobrychis (Fig. 69) ， and Sulla (Fig. tion may be a diagnostic character between 70) 70) of tribe Hedysareae were considered to Carmichaelieae and Clianthus. compose a monophyletic group ， Hedysaroid In Genisteae ，CAs were obvious1y divided cI ade ，from molecular data (Sanderson and into two types ，i. e. ， (1) oval and basal in Wojciechowski 1996). In the clade ，CAs Lupinus (Figs. 101 ，102) ， and (2) liuear and were situated at basal part (Fig. 69) in apical in Cytisus (Fig. 100) and Spartium Onobrychis ，basal-middle part (Figs. 47 ，64- (Fig. 103). In the tribe ， the shapes and po- 68) 68) in Alhagi and Hedys αrum ， or apical- sitions of CAs may be useful to consider middle middle part (Fig. 70) in Sulla ， of cotyledon inter-generic phylogenetic relationships. blades. blades. Sulla coronaria had been treated as

Hedysarum coronarium L. However ，re 田 Conclusion cently ， H. coronarium was considered to be Shape and position of CAs usually are treated treated under genus Sulla Medik. from mor- similar among the different species in the phological phological and palynological data (Choi and same genus and these characters may be Ohashi Ohashi 1996 ，1998). The difference of CA's useful to consider inter-generic phylogenetic position position between Sulla coronaria (Fig. 70) relationships in Papilionoideae. However ， in and and the five Hedysarum species (Figs. 64- Desmodium and lndigofera ， the shape var- 68) supports this treatmen t. In the ied or the position changed are exceptionally

Hedysaroid Hedysaroid cI ade ， the positions of CAs may found in the same genus ， and these charac 聞 be be useful to consider inter-generic phyloge- ters need to be further investigated to clarify netic netic -r elationships. their systematic utility in these genera. Genera Genera Biserrula (Fig. 49) ， Colutea (Fig. 51) ，Lesserti α(Fig. 57) ，Oxytropis (Fig. 58) ， This study was partly supported by Sutherlandia Sutherlandia (Fig. 59) ，Swainsoni α(Fig. F吋iwara Natural History Foundation (to Y. 60) ，and Clianthus (Fig. 1 in Endo and E.) ， Grants-in-Aid No. 07041122 (1 995- Ohashi 1998a) of the tribe Galegeae and 1996) and No. 10041153 (1 998) from the tribe tribe Carmichaelieae (Figs. 61-63) were pre- Monbusho International Scientific Research sumed to be members of a monophyletic Program (to H. 0.) ， and the Japan Society group ， Astragalean clade (Sanderson and for the Promotion of Science and the Korean Liston Liston 1995 ， Sanderson and Wojciechowski Science and Engineering Foundation. 1996). 1996). Shapes of CAs of this clade were oblong oblong (Figs. 49 ，51 ，58 ， 61-63; Fig. 1 in References Endo and Ohashi 1998a) or linear (Figs. 57 ， Bruneau A. ，Doyle J. and Doyle J. L. 1994. Phylo- 59 ， 60). Positions of the CAs were basal- genetic relationships in Phaseoleae: evidence from chloroplast chloroplast DNA restriction site characters. In: middle middle (Fig. 58) ， middle (Figs. 49 ，57 ，61- Crisp M. and Doyle J. (eds.) ，Advances in le- 63) 63) or apical-middle part (Figs. 51 ，59 ， 60; gume systematics 7: Phylogeny. pp. 309-330. Fig. Fig. 1 in Endo and Ohashi 1998a) of coty- Roya1 Botanic Gardens ，Kew. ledons. ledons. In the clade ，Carmichaelia and Choi B. H. and Ohashi H. 1996. Pollen morphology Clianthus Clianthus were considered to be phyloge- and of Hedysarum and its related gen- era era of the tribe Hedysareae (Leguminosae- netically netically cI osely related to each other from Papilionoideae). Papilionoideae). J. Jpn. Bo t. 71: 191-213. molecular data (Sanderson and 一一- and 一一一 1998. (1 377) Proposal to conserve the Wojciechowski 1996). However ，CAs of name Hedysarum (Leguminosae: Papilionoideae) Carmichaelieae Carmichaelieae were situated at the middle with a conserved type. Taxon 47: 877. part part (Figs. 61-63) and that of Clianthus was Doyle J. J. and Doyle J. L. 1993. Chloroplast DNA at at the apical-middle part (Fig. 1 in Endo and phylogeny of the Papilionoid 1egume tribe Phaseoleae. Phaseoleae. Systematic botany 18: 309-327. Ohashi Ohashi 1998a). The difference of ~he posi- 306 植物研究雑誌第74 巻第5号 平成11 年10 月

一一，一一， Ballenger J. A. ， Dickson E. E. ，K 吋ita T. 1-7. and and Ohashi H. 1997. A phylogeny of the chloro- Polhill R. M. 1994. Complete synopsis of legume plast plast gene RBCL in the Leguminosae: taxonomic genera. In: ILDIS and CHCD (eds.). Phytochemi- correlations correlations and insights into the evolution of cal dictionary of the Leguminosae ，vo l. 1. nodulation. nodulation. Am. J. Bo t. 84: 541-554. and their constituents. pp. xlix-lvii. Chapman & Endo Y. and Ohashi H. 1997. Cladistic analysis of Hall ， London. phylogenetic phylogenetic relationships among tribes Cicereae ， Sanderson M. J. and Li ston A. 1995. Molecular phy- Trifolieae ， and Vicieae (Leguminosae). Am. J. Bo t. logenetic systematics of Galegeae ， with special 84: 84: 523-529. reference to Astragalus. In: Crisp 乱1. and Doyle J. 一一 and 一一一 1998a. The features of cotyledon ar- J. (eds.) ， Advances in legume systematics 7: Phy- eoles eoles in Leguminosae and their systematic utility. logeny. pp. 331-350. Royal Botan Ic Gardens ，Kew. Am. J. Bo t. 85: 753-759. 一一一 and Wojciechowski M. F. 1995. Molecular phy- 一一- and 一一一 1998b. Variations in the anatomical logenetic analysis of a temperate legume clade features features of cotyledon areoles in Leguminosae. J. (). Am. J. Bot. (Supplement) 82(6): 159. Jpn. Jpn. Bo t. 73: 264-269. 一一-and 一一一 1996. Diversification rates in a tem- 一一-and 一一一 1999. The developmental change of cell perate legume clade: are there “ so many species" structures structures stained by iron-hematoxylin in the cot- of Astrag α lus (Fabaceae)? Am. J. Bo t. 83: 1488- yledon yledon areole of Leguminosae. J. Jpn. Bot. 74: 1502.

遠藤泰彦 a，大橋広好 b .マメ科マメ亜科における cotyledon cotyledon areole の形と位置の変異と分布，および その分類学的意義 マメ科マメ亜科の成熟種子内の子葉には表皮細 子葉の基部，中央基部寄り，中央，中央先端寄り， 胞がいぼ状に隆起する cotyledon areole (;;: CA) と 先端と様々である.これら外形および出現位置に いう構造が見られる (Endo and Ohashi 1998a). 本 ついては，一般に同一属内の種類の間では互いに 論文では， 23 連 98 属 147 種について CA の有無を 類似している.このことから，属間の系統関係を考 調査し(うち 109 種については Endo and Ohashi 察する際に有用であると考えられる.ただし，ヌス 1998a から引用) ， そのうち 22 連 69 属 108 種につ ピトハギ属，コマツナギ属では，同一属内で変異が いて CA の存在を確認した.この 108 種について 認められた.これら属においては，属内での変異の CA の外形および出現位置を図示した(うち 1種は 実態を明らかにし，その分類学的意義をより詳細 Endo and Ohashi 1997. 4 種は同 1998a から引用) . に検討する必要がある. さらに，その分類学上の有用性について議論を c千葉県立中央博物館植物学研究科 行った. CA の外形は三角形，円形，だ円形，長だ b 東北大学大学院理学研究科生物学教室) 円形，線形と様々な形態をとり，また，出現位置も