植物研究雑誌 J. J. Jpn. Bo t. 78:269-294(2003) 78:269-294(2003)

A New Circumscription of and a New Akschindlium (Leguminosae)

Hiroyoshi Hiroyoshi OHASHI

Botanical Botanical Garden ，Graduate School of Science ，Tohoku University ，Sendai ，980-0862 JAPAN (Received (Received on April 26 ， 2003)

The genus Tadehagi founded by the author in 1973 is revised. It is divided into two genera: genera: Tadeh α gi with a new circumscription and a new genus Akschindlium H.Ohashi based based on Tadehagi godefroyanum (Kuntze) H.Ohashi. Pollen features of three species representing representing the genera Tadehagi ， Akschindlium and 訂 e provided. The newly newly circumscribed Tadehagi is divided into two subgenera: Tadehagi and Kerria. A key key to these genera and subgenera is provided. Tadehagi comprises six species: T. alatum (DC.) (DC.) H.Ohashi ，T. auriculatum (DC.) H.Ohashi ，T. andamanicum (Balakr. & N.G.Nair) S.P.Mathew ，T. robustum Pedley ，T. rodgeri (Schind l.) H.Ohashi and T. triquetrum (L.) H.Ohashi. H.Ohashi. varies in the leaf shape and loment hairiness especially in in the Lesser Sunda Islands ，New Guinea and the Philippines. The species is newly cir- cumscribed cumscribed as including subsp. pseudotriquetrum (DC.) H.Ohashi and a hybrid between subsp. subsp. pseudotriquetrum (DC.) H.Ohashi and subsp. triquetrum ， nothosubsp. philippinensis philippinensis H.Ohashi. The hybrid includes a variety ，var. palawanensis H.Ohashi. Tadehagi Tadehagi triquetrum subsp. triquetrum is newly recorded in Taiwan. A key to these spe- cies ，subspecies and hybrids is also provided.

Key words: Akschindlium ， ，Droogmansia ，Leguminosae ， Tadehagi.

The genus Tadehagi H.Ohashi was estab- calyx ， seeds and bracteoles ， but some charac- lished lished in 1973 based on Desv. ex ters are close to Tadehagi such as the smaller Benth. ， a later homonym of Pteroloma flowers ， narrowly oblong loments with less Hochs t. & Steud. ，and a species of Droog- constricted sutures ， narrowly winged petiole ， mansia De Wild. (Ohashi 1973). It included and pollen morphology. three three subgenera ，i. e. ，Tadehagi s. s甘.， Kerria Pollen grains of Godefroya ，Tadehagi and H.Ohashi ， and Godefroya (Schind l.) H. Droogmansia are fundamentally similar in Ohashi. Ohashi. Tadeh α gi was ，however ， recognized morphology as well as wall stratification. as as intergrade with Droogmansia (Ohashi This overall similarity indicates their close et et al. 1981). This view has been accepted relationship ， but the tectum differs between due to the problematical taxonomic position Droogm α nsia and the other two (Ohashi of of Tadehagi godefroyanum (Kuntze) H. 1971). Godefroya and Tadehagi have reticu- Ohashi ， the monotypic representative of the late to foveolate sculpturing at the subgenus Godefroya (Fig. 1). mesocolpium with the lumina 0.5-1.5 flIIl in Subgenus Godefroya was founded by diameter ， while Droogmansia has reticulate Schindler Schindler (1924) in the genus Droogmansia. sculpturing with lumina 1-3 戸n in diameter It It is similar to Droogmansia in the stamens ， (Figs. 2-4).

-269- 270 270 植物研究雑誌第78 巻第5号 平成15 年10 月

In In addition to the similarities in macro- found entirely inside the East Asian distri- scopic scopic and pollen characters between bution range of Tadehagi. Droogmansia is Godefroya Godefroya and Tadehagi ， Godefroya is confined to tropical Africa. Therefore it was

Fig. Fig. 1. Akschindlium god ψ， 'oyanum (Kuntze) H.Ohashi. Thailand. Smitinand 10060 (BKF ，cited in Ohashi 1973). October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 271

natural natural to accommodate Godefroya within in any other part of the world (Ohashi 1973 ， Tadehagi Tadehagi (Ohashi 1973). Reassessment of 1992). The genus is consid- the the morphological and pollen characters as ered to be most primitive genus among outlined outlined above suggested that Godefroya is and its related genera and apparently apparently intermediate between Tadehagi Tadehagi ， in the older sense including and and Droogmansia ， in spite of the shared dis- Godefroya ，was supposed to be derived from tribution tribution of the former two genera. Tadehagi Dendrolobium (Ohashi 1973). Dendrolobium and and Droogmansia are recognized as distinct is most abundant in Indochina with several in in results of an analysis of the chloroplast endemic species. Akschindlium and Tade- gene gene rbcL (Bailey et al. 1997 ， Kajita et al. hagi subgenus Kerria are distributed in simi- unpublished) ， although Godefroya is not yet lar areas; the former is restricted to Thailand analyzed. analyzed. The chromosome numbers are and Indochina (Cambodia ， Laos and known only in Tadeh α gi triquetrum subsp. Vietnam) ， and the latter Myanmar ， Thailand triquetrum triquetrum and subsp. pseudotriquetrum ， and Indochina (Laos). Subgenus Tadehagi is i. i. e. ，n = 11 and 2n = 22 (cf. Ohashi 1973 ， widely distributed in SE. and E. Asia and N. Goldblatt Goldblatt and Johnson 1991 ， 1998). Al- Australia ， while Droogmansia is endemic to though though knowledge of these taxa 訂 e insuffi- tropical Africa. It may be supposed that cient cient at the moment ， I would favour Akschindlium and Tadehagi subgenus Kerria recognition recognition of the subgenus Godefroya as an originated in Indochina from a species of independent independent genus distinct from Tadehagi Dendrolobium and that Akschindlium might and and Droogmansi αfor stability of the system be phylogenetically distinct from Droog- of of tribe Desmodieae. To combine these three m αnsza. taxa taxa into one genus would be a revival of the This work had been prepared mainly by wider wider concept of Desmodium proposed by studying herbarium specimens of Tadehagi Bentham (1 865) that is inconsistent in its and its related genera kept in A & GH ，BKF ，

boundary boundary with ， ， or Alysi 国 BO ，CAL ，K ，KYO ，L ，LAE ，NY ，PNH ，TI ， cα rpus. I will discuss the problems in generic TUS ， and US. Studies on the species from circumscription circumscription in the tribe Desmodieae in a Thailand (Ohashi 1982) ， Indo-China (Dy precursory precursory treatment for Desmodium and its Phon et al. 1994) and Malesia (Ohashi in allied allied genera in Malesia (Ohashi in prepara- prep.) were especially helpful for this revi- tion). tion). sion. Field observations were made in The new generic name for Godefroya ， Australia ， China ， the Philippines and Akschindlium Akschindlium H.Ohashi ，is proposed here in Taiwan. Results of pollen mo 叩hology de- honor honor of Dr. Anton Karl Schindler (1 879- scribed in this paper were provided by Ye. 1964) 1964) who founded subgenus Godefroya in Pollen materials were obtained from herbar- Droogmansia and ，also ，made great contribu- ium specimens. Methods used for pollen ex- tions tions to the of the tribe amination and terminology for pollen Desmodieae in Leguminosae (see Schubert descriptions in this study 紅 e the same as 1964 ， Pedley 1999). those used in Ye and Ohashi (2002). The tribe Desmodieae is most diverse in the the Sino-Indian region (Ohashi et al. 1981). Pollen morphology Desmodium and its related genera 紅 e con- Pollen grains of Akschindlium ，Droog- sidered sidered to have undergone their prim 紅 y ra- mansia ， and Tadehagi were first described diation diation in SE. Asia because more high level by Ohashi (1 971 ， 1973) using light micros- taxa taxa (genera ， subgenera and sections) of copy. Descriptions and figures 紅 e provided Desmodium and allies occur in this 紅 ea than here from results obtained by scanning elec- 272 植物研究雑誌第78 巻第5号 平成15 年10 月 tron tron microscopy (SEM) and transmission General features of pollen grains of electron electron microscopy (TEM). SEM and TEM Akschindlium ，Droogmansia ， and Tadehagi

pollen pollen features in these genera were first (Figs. 2 -4): The pollen grains are tricol 回 shown in Droogmansia scaettaiana by porate; colpi 0.8-0.9 times and endoaperture Ferguson Ferguson and Skvarla (1 981). 0.2-0 .3 times the length of polar axis; exine

Fig. Fig. 2. Pollen morphology of Akschindlium god そ斤 oyanum (Schind l.) H.Ohashi. A: Equatorial view. B: Tectum at mesocolpium. C: Equatorial view. D: view. Polar E: Cross section at mesocolpium. F: Cross section. section. G ， H: Cross sections at aperture. Scale bars: A ，C ，D ，F = 5 Il m ，B ，E ，G ，H = 1μm. Voucher specimen: specimen: G. Murata & al. T-37621 (TUS). October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 273

0.8- 1. 5 阿n thick at mesocolpium; tectum ， H.Ohashi (Fig. 2): Pollen grains tricolporate ， infratectum infratectum with foot layer and endexine al- (20.8 一)22.5(-25.6)μm in polar axis ，(2 1. 3一) most most equal in thickness ， each about 1/ 3 of 22.9(-25.0) 伊n in equatorial diameter; P厄= the the exine; columellae regul 訂; foot-layer con- (0.91-) 0.98 (-1. 05) ， oblate spheroidal to tI nuous. prolate spheroidal; colpi broad ， ca. 4 阿n wide wide at equator ，tapering to pointed ends ， Akschindlium Akschindlium god ψoyanum (Kuntze) colpus colpus membrane almost smooth ，occasion-

Fig. Fig. 3. Pollen mo 中hology of Droogmansia pteropus (Baker) De Wild. A: Equatorial view. B: Tectum at mesocolpium. mesocolpium. C: Equatorial view. D: Polar view. E: Cross section at mesocolpium. F: Cross section. G: Cross Cross section at aperture. Scale bars: A ，C ，D ，F = 5 ドm ，B ，E ，G ，H = 111m. Voucher specimen: Stolz 2384 (A). 274 274 植物研究雑誌第78 巻第5号 平成15 年 10 月 ally ally with fine granules ， margins almost un- Droogm αnsi α pteropus (Baker) De Wild. differentiated differentiated from mesocolpium; sexine (Fig. 3): Pollen grains tricolporate ，(2 1. 6-) tectate; tectate; sculpture at mesocolpium reticulate 25.3(-27. 1)阿 n in polar axis ，(1 9.5-)2 1. 7 to to foveolate ，occasionally with perforate ， (-23.2) 阿n in equatorial diameter; P厄= lumina lumina 0.5-1.2 阿n in diameter. Exine ca. 1.5 (1. 08-) 1. 17(- 1. 27) ， prolate spheroidal to 阿n thick in mesocolpium ， sexine about as subprolate; colpi broad ， ca. 4 阿n wide at thick thick as nexine; tectum discontinuous ， about equator ， tapering to rounded ends ， colpus 1/ 3-114 times the thick of exine; infratectum membrane almost smooth ，occasionally with well well defined ， as thick as tectum ，consisting fine granules ， margins with a punctate; of of regular columellae; foot-layer continuous ， sexine semi-tectate; sculpture at mesocol- less less than 1/ 2 the thickness of infratectum ， pium reticulate ， lumina 1-3 阿n in diameter ， consistent; consistent; endexine well developed ， about with columellae elements. Exine ca. 1.5 阿n 1/ 3 the thickness of exine ，gradually thicken- thick in mesocolpium ， sexine slightly thicker ing ing toward colpi. than nexine; tectum discontinuous ， about 1/ 3 Voucher Voucher specimen: Thailand. E; Surin Prov. G. the thickness of exine; infratectum about 1/ 2 Murata Murata & al. T-37621 (TUS). the thickness of tectum thickness ，consisting

Fig. Fig. 4. Pollen morphology of Tadehagi triquetrum (L.) H.Ohashi subsp. triquetrum. .A: Equatorial view. B: Tectum at mesocolpium. C: Equatorial view. D: Polar view. E: Cross section at mesocolpium. Scale bars: bars: A ，C ，D = 5μm ，B ，E= 1μm. Voucher specimen: Y. Endo 2 Dec. 1981 (TUS). October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 275 of of columellae; foot-layer continuous ，same ...... Droogmansia thickness thickness as columellae ，consistent; endexine 2. Free parts of filaments elliptic ，imbricate. developed ， about 1/ 4 the thickness of exine ， Wings larger than the keels. Loments gradually gradually thickening toward colpi. short stalked; articles longer than broad. Voucher specimen: Kyimbila. Stolz 2384 (A). Stipules connate into a bifid scale ， early deciduous. deciduous. Endemic to Thailand and Tadeh α gi triquetrum (L.) H.Ohashi subsp.

Indo-China Indo-China ....・ H ・...・ H ・...・ H ・... Akschindlium triquetrum triquetrum (Fig. 4): Pollen grains tricolpo- 1. 1. Free p紅白 of filaments uniform ，filiform; rate ，(1 6.9-) 20.1 (-24.0)μm in polar axis ， stipules stipules free ，bracteoles present; lowest (= (13.3 ー)17.2(-24 .4 )μm in equatorial diame- abaxial) abaxial) calyx-lobes longer than upper and ter; ter; PA = (0.94 ー) 1. 19 (-1. 72) ， oblate sphe- lateral lateral lobes; seeds rim-arillate around the roidal ，prolate spheroidal ，subprolate prolate; hilum...... Tadehagi hilum...... Tadehagi colpi colpi n紅 row ， ca. 1μm wide at equator ， 2. 2. Wings almost equal to the keel-petals in tapering tapering slightly to rounded ends ， colpus size size and both petals not arcuate; pods membrane with fine granules ， margins with long-stiped long-stiped (the stipe more than 8 mm a complete tectum; sexine tectate; sculpture long) ，distinctly constricted along the at at mesocolpium reticulate to foveolate ，occa- lower lower margin; petiolar wings inconspicu- sionally sionally with perforate ， lumina 0.8-1.5μmin ous. ous. Endemic to Myanmar ， Thailand and diameter. diameter. Exine 1- 1. 5 阿n thick in meso-

Laos.... ・H ・....・ H ・....・ H ・....・ H ・...subgen. Kerria colpium ， sexine thicker than nexine; tectum 2. 2. Wings distinctly longer than the keel- discontinuous ， about 1/ 3 the thickness of petals petals and both petals apparently arcuate; exine; exine; infratectum well defined thickness pods pods shortly stiped (the stipe less than 7 same as tectum ， consisting of regul 訂 mm long) ， shallowly constricted along columellae; columellae; foot-layer continuous ， less than the the lower margin; petiolar wings con- 1/ 3 the thickness of infratectum; endexine spicuous. spicuous. Tropical Asia ， N. Australia and well well developed ， about 1/ 3 the thickness of

Pacific... ・H ・..…...・ H ・..…… subgen. Tadehagi eXlne. eXlne. Voucher specimen: Cultivated in Tohoku 1. 1. Akschindlium H.Ohashi ， gen. nov. University ， Sendai. Originally seeds 合omHongkong. Y. Y. Endo 2 Dec. 1981 (TUS). Droogm α nsia subgen. Godefroya Schind l. in in Repert. Spec. Nov. Regni Veg. 20: 274 Taxonomic treatment (1 924) [Type species: Droogmansia godej シ0 ・ Key to Akschindlium ，Droogmansia and yana (Kuntze) Schind l. (= Meibomia gode- Tadehagi Tadehagi with its subgenera: froyana Kuntze)]. 1. 1. Free p紅白 of filaments dimorphic ，alter- Tadehagi subgen. Godefroya (Schind l.) nately nately expanded or filiform; bracteoles ab- H.Ohashi in Ginkgoana 1: 284 ，t. 44a (1 973). sent; sent; upper (= adaxial) calyx-lobes longer Meibomia auc t. non Heis t. ex Fabr.: than than lateral and lowest lobes; seeds Gagnep. in Fl. Indo-Chine 2: 608 (1920) ， ut slightly slightly or sc 訂 cely rim-arillate around the “Meibomi αAdanson". hilum...... 2 hilum...... 2 Frutex ， erectus ， ramosus; rami teretes; 2. 2. Free p紅 ts of filaments narrowly folia stipulata ， petiolata ， 1- foliolata ， 2- obcordate ， not overlapping each othe r. stipellata; petiolo alato; stipulae connatae ， Wings smaller than the keels. Loments bifidae ， caducae; inflorescentiae terminales usually usually long stalked; articles as long as et axillares ， racemosae; flores in bractearum or or longer than broad. Stipules free ，per- axillis ， 2-5 in quoque fasciculo ，pedicellati; sisten t. Endemic to 佐opical Africa.... bracteae primariae et secundariae praesentes; bracteolae bracteolae nullae; calyx late campanulatus ， 276 276 植物研究雑誌第78 巻第5号 平成15 年10 月

4-fidus ， lobus posticus magnissimus; (1 920). vexillum vexillum orbiculare vel perlate obovatae ， Droogmansia go d， ψoyana (Kuntze) exauriculatum; exauriculatum; alae carinam superantae; Schind 1. in Reper t. Spec. Nov. Regni Veg. stamina stamina monadelpha ， vexillari 2/3 longi- 20: 274 (1 924) ， in Repert. Spec. Nov. Regni tudinis tudinis cum ceteris connato ，libero ad apicem Veg. Beih. 49: 310 (1 928); Thuan ， F l. Camb. et et basim ， filamentis connati libero ad Laos Vietn. 23: 126 ，t. 21 (1 987); Lock &

apiceum ， filamentorum dilatatorum partes Heald ， Leg. Indo-China Check 四 Lis t: 75 liberae liberae latae ， ovatae et ellipticae ，imbricatae; (1 994). ovarium ovarium breviter stipitatum; disco tubuli- Desmodium god ψ， oyanum (Kuntze) formi formi circumcinctum; stigma terminak; Craib ， F l. Siam. Enum. 1: 407 (1 928); Me 町. legumen legumen stipitatum ，compressum ，angusto- in Brittonia 2: 193 (1936).

oblongum ，articulatum; semina rim-arillata. Tadehagi god ψ， oyanum (Kuntze) Shrub ， erect ， usually 1. 5-3 m high; H.Ohashi in Ginkgoana 1: 284 ， p l. 44a ，figs. branches branches terete. Leaves 1- foliolate. Petioles 81-83 (1 973); Dy Phon in Dy Phon et al.， Fl. winged. winged. Stipules connate at the opposite side Camb. Laos Vietn. 27: 48 (1 994). of of the petioles ，bifid at apex ， early decidu- Distr.: Thailand and Indo-China ous ， leaving a transversely long sc 紅 on the (Cambodia ，Laos ， and Vietnam). branch. branch. Inflorescences pseudoracemose ，ter- Specimens. examined. (addition to Ohashi 1973): minal minal and axillary. Flowers in 2-5-flowered Thailand. Surin. G. Murata et al. T-37621 (KYO ， fascicles. fascicles. Primary and secondary bracts pre- TUS). Cambodia. Mout Kulen. Martin 1275 (P ， TUS). Laos. Laos. Vientiane. Hayata 8 Dec. 1921 (TI ， TUS); Dan sen t. Bracteole absen t. Calyx broadly Si. Si. Iwatsuki et al. 1043 (TUS); M t. Phu Kao Kwai. campanulate ，4-10bed; upper lobe larger than Iwatsuki et al. 1012 (TUS); Se Ban F剖. Poilane 28277 other other lobes. Standard orbicular or very (P ， TUS). broadly broadly obovate ， not auriculate; wings larger The stamen of Akschindlium god ψ， 'oya- than than the keel-petals. Stamens monadelphous ， num is monadelphous ， but its vexi l1 ary sta- vexillary vexillary one free at the apex and base and men is free at the very base and has sma l1 connate connate with the others below two thirds of openings (fenestrae). It has a disk around the its its length; connate stamens free at the apex ， base of ov 紅 y. These features in the structure free free part of the filaments much dilated ，over- of flowers indicate that the genus is adapted lapping lapping each other ，membranous ， e l1 iptic. to bee pollination. Ovary shortly stiped. Disk distinct around the the base of ovary. Loments short1y stiped ， 2. Tadehagi H.Ohashi ， Ginkgoana 1: 280 compressed ，straight ， narrowly oblong; arti- (1973) ，p. ，exc l. subgen. Godefroya [Type: cles cles quadrate to transversely oblong ，reticu- T. triquetrum (L.) H.Ohashi (= Hedysarum late-veined. late-veined. Seeds scarcely rim-arillate triquetrum L.)]; H.Ohashi & al.， Adv. Leg. around around the hilum. Sys t. 1: 299 (1 981); Huang & H.Ohashi ， F l.

Type species: Akschindlium god ザヤ， oya 幽 Taiwan ed. 2 ，3: 350 (1 993); Dy Phon in Dy num (Kuntze) H.Ohashi ( = Meibomia gode- Phon et al.， Fl. Camb. Laos Vietn. 27: 47 斤oyana Kuntze). (1 994); Yang & Huang in Fl. Reipub. Popu l. Sin. Sin. 41: 62 (1 995); Pedley ， Rev. Hand. Fl. 1-1. 1-1. Akschindlium godefroyanum Ceylon 10:168 (1 996); H.Ohashi in Sci. Rep. (Kuntze) (Kuntze) H.Ohashi ， comb. nov. (Fig. 1). Tohoku Univ. ser. 4，Bio l. 40: 249 (1 999);

Meibomia god げ示。， yana Kuntze ， Rev. Gen. Pedley in Austrobaileya 5: 258 (1 999). 1: 1: 196 (1 891) [Type: Cambodia. Godefroy Desmodium sec t. Pleurolobium DC. [un- s.n. s.n. July 1875. (P holo. ，iso. ，n. v.)]; Gagnep. ranked] Pteropoda DC. ，Prodr. 2: 326 (1 825) in in Lecomte ， F 1. Indo-Chine 2: 608 ，fig. 56 [Type species (here designated): Desmodium October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 277 triquetrum triquetrum (L.) DC.] & Mem. Leg. 317 to deeply constricted. Seeds transversely (1 826). Ohashi (1 973) regarded Pteropoda broadly to transversely elliptic ，rim-arillate ， DC. at the subsection ran k. concave at the hilum. Pteroloma Pteroloma Desv. ex Benth. in Miq. ， P l. Jungh. Jungh. 217 ， in adnota ，& 219 (1 852) [Type 2-1. Tadehagisubgen. J( er ・ria H.Ohashi in species: species: P. triquetrum (L.) Desv. ex Benth. Ginkgoana 1: 287 (1 973) [Type species: T. (=Hedys αrum triquetrum L.)] ，non Ptero- kerrii (Schind l.) H.Ohashi (= Pteroloma loma Hochs t. & Steud. (1 841); Schind l. in kerrii Schind l.)]. Repert. Repert. Spec. Nov. Regni Veg. 20: 271 (1 924) ， in Repert. Spec. Nov. Regni Veg. 2-1- 1. Tadehagi rodgeri (Schind l.) Beih. Beih. 49: 362 (1 928); Hutch. ， Gen. Fl. Pl. 1: H.Ohashi in J. Jpn. Bo t. 57: 266 ，fig. 1 484 (1 964) ，ut “ Pteroloma DC."; Allen & (1 982). (Fig. 5) Allen ， Leguminosae: 568 (1 981). Pteroloma rodgeri Schind l. in Repert. Desmodium sec t. Pteroloma (Desv. ex Spec. Nov. Regni Veg. 20: 272 (1 924) Benth.) Benth.) Benth. in Benth. & Hook .f.， Gen. Pl. [Type: Burma (Myanmar). Ruby Mines ， 1: 1: 519 (1865); Merr. in Philipp. 1. Sci. ， C. Mogo k. A. Rodger 196 (K holo. ，CAL iso.)]. 5: 5: 86 (1910). Pteroloma kerrii Schind l. in Repert. Spec. Desmodium subgen. Pteroloma (Desv. ex Nov. Regni Veg. 23: 361 (1 927) [Type: Be 印nt 血h.) Baker 泊in Ho ∞依k.工，O Fl. Bri t. Ind. 2: Thailand. Me Ping Rapids ，Keng Soi ， al t. ca. 163 163 (1 876). 400 m. F. Kerr 4634 (K holo.)]. Meibomia Adanson (1 763) sensu Gagnep. Desmodium kerrii (Schind l.) Craib ， F l. in in Lecomte ， F l. Indo-Chine 2: 608 (1 920). Siam. Enum. 1: 410 (1 928). Subshrubs Subshrubs or shrubs. Leaves l-foliolate. Pteroloma triquetrum subsp. rodgeri Stipules Stipules free. Petioles winged. Stipels 2，con- (Schind l.) H.Ohashi in J. Jpn. Bo t. 41: 96 nate nate with the petiolar wings. In f1 0rescences (1 966). pseudoracemose ， terminal or axillary ，usu- Tadehagi triquetrum (L.) H.Ohashi subsp. ally ally 2-6- f1 owered at a node. Primary bracts rodgeri (Schind l.) H.Ohashi in Ginkgoana 1: be 紅 ing 2 second 紅 y bracts; secondary bracts 295 (1 973); Dy Phon in Dy Phon & al.， Fl. bearing bearing f1 0wers at the axi l. Bracteoles pre- Camb. Laos Vietn. 27: 54 ， p l. 11: 6-10 sen t. Calyx campanulate ，4-10bed; upper lobe (1 994). re f1 exed ， broadly ovate-triangular ，entire or Tadehagi kerrii (Schind l.) H.Ohashi in minutely minutely 2-toothed at the apex ， lowest lobe Ginkgoana 1: 287 ， p l. 43 (1 973). longer longer than other lobes. Distr.: 民1yanmar ， Thailand and Laos. Petals Petals with well developed nerves; stan- Specimens examined (exc l. ci t. in Ohashi 1973 ， dard dard clawed ，re f1 exed at the top of the claw ， 1982). Myanmar. Ruby Mines Distric t: Mogok ， blade blade orbicular to transversely broadly ellip- Shweli Valley. A. Rodger 196 (K holo. of Pteroloma tic tic or obovate; wings elliptic oblong ， not rodgeri Schind l.， CAL iso. The type in CAL is isotype ， not not as in Ohashi 1973). Thailand. 乱1e Ping Rapids ， shorter shorter than the keel-petals. Stamens 10 ， Keng Soi ， al t. ca. 400 m. F. Kerr 4634 (K holo. of monadelphous ，vexillary stamen free at the Pteroloma kerrii Schind l.); loc. ci t. Kerr 4641 (BK ， not apex ， connate to others two thirds of its the type ， not as in Ohashi 1973); C. F. van Beusekom length. length. Disks presen t. Ovary stipitate ，5-8- & Phengklai 2569 (L); Prov. E. ， Chaiyaphum. Nam 0 ovuled; ovuled; style glabrous; stigma capitate. Pham. 16 2WN 101 0 45 包 al t. 600 m. C. F. van Beusekom & al. 4138 (KYO). Loments stipitate ， straight or slightly incurved ，compressed ， narrowly oblong ，usu- 2-2. 2-2. Tadeh α gi subgen. Tadehagi: H. ally ally 5-8 jointed ， upper sutures nearly straight Ohashi Ohashi in Ginkgoana 1: 290 (1 973) [Type or or slightly undulate ， lower sutures shallowly 278 植物研究雑誌第78 巻第5号 平成15 年10 月 species: species: T. tr 匂uetrum (L.) H.Ohashi (= Key to the species of subgenus Tadehagi: Hedysarum tr 匂uetrum L.)]. 1. Loments distinctly stiped ， exserted from

the the calyx; stipe 3-7 mm long ...・ H ・-… H ・H ・.2

Fig. Fig. 5. Tadehagi rodgeri (Schind l.) H.Ohashi. Thailand. Prov. E. ，Chaiyaphum. Nam Pham. C. C. F. van Beusekom & al. 4138 (KYO). October October 2003 Joumal of Japanese Botany Vo l. 78 No. 5 279

2. 2. Loments 2-seeded ， lateral surfaces subsp.α latum (DC.) H.Ohashi in J. Jpn. Bo t.

puberulent; puberulent; articles 13-16 mm long.... ・H ・ 41: 95 (1 966).

...・ H ・..… .T. andamanicum Tadehagi tl 匂uetrum subsp. alatum (DC.)

2. 2. Loments more than 4オointed ，lateral sur 国 H.Ohashi in Ginkgoana 1: 292 ，t. 41a (1973). faces faces glabrous or glabrescent; articles Shrub. Leaves l-foliolate. Stipules nar-

less less than 10 mm long.... ・H ・....・ H ・-…...・ H ・..3 rowly ov ate ， 1.5-2 cm long ， 3-5 mm wide ， 3. 3. Pericarp thinly chartaceous ， margins acuminate ，glabrous ，conspicuously striate. pubescent; pubescent; articles broadly oblong ， 6-9 Petioles 1-3 cm long ，winged ， sp 紅 sely mm long ， 7-12 mm wide.... … T. alatum appressed pubescent; wings 2-4 mm wide at 3. 3. Pericarp corky ， margins glabrescent ， broadest part ， almost glabrous; joined with stipes stipes 3-4 mm long; articles almost stipel at tip; stipel curved ，glabrous. Leaflet- quadrangular ， 5-7 mm long and wide... blade narrowly ovate or narrowly elliptic ，7- .T. ….T. auriculatum 15.5 cm long ，1. 5-3 cm wide ， acuminate at 1. 1. Loments shortly stiped ， not or sc 紅白ly apex ， obtuse to rounded or sometimes exserted exserted from the calyx; stipe 1-3 mm slightly cordate at base ， both surfaces long; long; articles in middle part less than 5 sparsely appressed pubescent on midrib ， 訂 訂 mm long.... ・・ H ・H ・.....・ H ・...・ H ・-…...・ H ・....・ H ・-….4 sp sely appressed ciliate along m gin ， 4. 4. Loments 4-勾 ointed ，sp 紅 sely hairy seemingly glabrous but minute hairy in other on sutures; articles ca. 5 mm long ，8 parts ，lateral nerves alternate ，1 4- 18 pairs ， mm wide; leaflet-blade less than 3 looped within m 訂 gin. times times longer than broad ， oblong or Inflorescences pseudoracemose ， to 40 cm

ovate...... ・H ・.....・ H ・.....・ H ・..… .T. robustum long ， 2-6 flowers per node ，rachis sp 紅 sely 4. 4. Loments 5-8-jointed ， densely hairy appressed pubescent ， with patent minute on sutures; articles 2.5-3.5 mm long ， hooked hairs. Primary bracts narrowly ovate 5-6 5-6 mm wide; leaflet-blade usually or narrowly triangular ， 3-5 mm long ，ca. 0.7 more than 3 times longer than broad ， mm wide ，striate ，ciliate; secondary bracts linear linear to narrowly oblong-ovate or nar- subulate ， 1-2 mm long. Pedicels 5-7 mm

rowly rowly ovate. …....・ H ・....・ H ・.T. triquetrum long ， covered sparsely to uniformly with pat- ent ent minute hooked hairs. Bracteoles subu- 2-2- 1. Tadehagi alatum (DC.) H.Ohashi ， late ， 1- 1. 5 mm long. Calyx appressed sta t. nov. (Fig. 6). pubescent and minute hooked hairy; lateral Desmodium alatum DC. ， Prodr. 2: 326 lobes triangular ， 1-1.5 mm long; the lowest (1 825) [Type: India orien t. ad Cawnpore (G lobe narrowly triangular ，3.5 -4 mm long. holo. ， n. v.); in Candolle: Prodromi Corolla pink to purplish red; standard Herbarium ，IDC micro-edition (TUS)] & incurved ， wings 5-6 mm long ， as long as the Mem. Leg.: 321 (1 826). keel ， long clawed (claw ca. 2 mm long) ， keel Desmodium triquetrum (L.) DC. subsp. distinctly auriculate ， long clawed (claw ca. 2 alatum alatum (DC.) Prain in 1. Asia t. Soc. Bengal ， mm long). Ovary densely pubescent along t. Pt. 2，Na t. His t. 66: 390 (1 897); Meeuwen in both m 紅 gins ， minutely stipitate. Reinwardtia Reinwardtia 6: 263 (1962); Sanjappa ， Leg. Loments 6-7 サointed ， dark red in young Ind. Ind. 164. (1992). and brown in mature ，stipitate (stipe 3-7 mm Pteroloma Pteroloma pseudotriquetrum xα latum: long) ，glabrous on lateral surfaces ，densely to Schind l. in Repert. Sp. Nov. 20: 272 (1 924). sparsely appressed or patent minute puber- [Voucher [Voucher specimen: India. Benga l. Regio ulent along both sutures ，reticulate on lateral tropica. tropica. Hooker f. & Thomson (K ， GH)]. surfaces ，pericarp thinly chartaceous; articles Pteroloma Pteroloma triquetrum (L.) Desv. ex Benth. broadly oblong ， 6-9 mm long ，7-12 mm 280 植物研究雑誌第78 巻第5号 平成15 年 10 月

Fig. Fig. 6. Tadehagi alatum (D C.) H.Ohashi. a: Bangladesh. Chittagong. Rich soil Ihandachery Hill40 miles miles from Chittagong. D r. Ki ng's collector Badul Kh an 207 (CAL ，cited in Ohashi 1973) ， b. Loments. Loments. Java. unknown collector ，Communica t. ex Herb. Lugd.-Ba t. (GH). October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 281 wide. wide. scrub forests on open hill slopes. 2 Dec. Distr.: Distr.: East India (Assam) and Bangladesh 1976. Balakrishnan & Nair 4798A (CAL (Chittagong) (Chittagong) . holo. ， n. v.)]. Specimens Specimens examined (exc 1. specim. cited in Ohashi Erect shrub ，1-2 m high. Leaflet-blades 1973). 1973). India. Bengal ， Regio trop. Hooker & Thomson oblong-elliptic ， 2.5-7 cm long ， 1-3.5 cm (GH ，L); Assam. Kh asia. regio 仕op. (K); Kh asia. C. B. wide. Pods stalked ，2-seeded ， 2.8-3.5 cm Clarke Clarke 14469C (K) ，C. B. Clarke 14476A (K) ，Monsto. long; long; articles 13-16 mm long ，9-10 mm 3000 3000 ft. C. B. Clarke 14480A (K); Kh asia. N. Cachar Hills ，at foot of landslide in full aftemoon sunshine ， wide ，puberulous ， finely reticulate. ground ground firly damp ， al t. 600 ft. Fl wrs pink; dark red Distr.: India. North Andaman Island. seed seed pods; lves light grey-green. Nov. 197 1. Yandell This is an insufficiently known species. I 50 50 (K ，L). Bangladesh. Chittagong ，Chundragona. 12 was unable to examine specimens of the spe- Jan. Jan. 1869. C. B. Clarke 8293 (K) ，Bindralun. 29 Feb. cies. Tadehagi triquetrum subsp. andama- 1868. 1868. C. B. Clarke 6579 (K). Chittagong Hil 1. Li ster in Bal 紘 r. & N.G.Nair was characterized 1876 1876 (L); Kaptai ，Chittagong Hill Tracts. Shrub 4 f t. nicum high. high. Fl s reddish. Parkinson 4284 (A ， GH); 5 km N. of in having 2-seeded pods of which the articles Teknaf. Teknaf. van der Maesen 3767 (K). Jawa (cultivated). are longer ，13-16 mm long ， than those in Beitenzorg. Beitenzorg. Erik Nyman (UPS); Ex Hort. Bogorensis ， other taxa (Balakrishnan and Nair 1982). Teysmann Teysmann 1860 (A ，BO ，K ，L); Cul t. in Hort. Bog. The subspecies appe 紅 s to belong to the sub- Halliez Halliez in anno 1893 (BO); unknown collector ， genus Tadehagi and close to T. auriculatum Communica t. ex Herb. Lugd.-Ba t. (GH ， K ， L). Madagascar Madagascar (introduced). Ch. D' Alleizette Nov. 1906 in floral characters. Sanjappa (1992) re 田 (L). (L). garded it as identical with Desmodium Schindler Schindler (1 924) regarded many speci- triquetrum subsp. auriculatum (DC.) Prain ， mens of a collection by Hooker f. and but the loments and leaflet-blades differ from Thomson in tropical Bengal as a hybrid be- the latter. Judging from the description and tween Pteroloma alatum and P. pseudo- the figure by Balakrishnan and Nair (1982) ， t1 匂uetrum ， because their mature loments the leaves are similar to those of Tadehagi were sterile~ Specimens of the collection I triquetrum subsp. pseudotriquetrum and the have so far examined are ，however ，referable pods 紅 e unique in the genus. The loment il- to to Tadehagi alatum in having a distinct stipe lustrated in Balakrishnan and Nair's Fig. 1 of the loment ， long pedicels ， long n紅 row (1982) has a distinct stalk which is exserted leaflets leaflets (the blade 14- 16.5 cm long and 3-4 from the calyx and seems to be more than 5 cm wide) ，and glabrous but ciliate pods. mm long ， although the loment appe 紅 s to be Tadehagi Tadehagi alatum always has ciliate pods ，al- immature because the stigma still remains on though it was characterized by Prain (1 897) the pods. Further studies on this species 紅 e as as glabrous throughou t. The hairiness on the needed based on ample materia l. loments loments in T. alatum shows the same pattem as as T. triquetrum subsp. pseudotriquetrum. 2-2-3. Tadehagi auriculatum (DC.) H. Schindler' Schindler' s presumed hybrid ，therefore ，ap- Ohashi ，sta t. nov. (Fig. 7). parently parently belongs to T. alatum. Desmodium auriculatum DC. ， Prodr. 2: 326 (1 825) [Type: Unknown locality but er- 2ふ 2. Tadehagi andamanicum (Bal 紘 r. roneously located as “Timor ぺwith a note as & N.G.Nair) S.P.Mathew in 1. Econ. Tax. “ Pteroloma auriculatum Desv!" Unknown Bo t. 14: 236 (1 990). collector (P holo. ， n. v.)] & Mem. Leg.: 321 Tadehagi Tadehagi triquetrum subsp.α ndamanicum (1826). Balakr. Balakr. & N.G.Nair in Bull. Bo t. Surv. India Desmodium tl 匂uetrum subsp. auricula- 24: 24: 29 ， fig. 1 (1 983) [Type: North Anda- tum (DC.) Prain in J. Asia t. Soc. Bengal ， P t. mans. Saddle Peak ，650-720 m above m.s. l.， 2，Na t. His t. 66: 390 (1897); Meeuwen in 282 282 植物研究雑誌第78 巻第5号 平成15 年10 月

Fig.7. Fig.7. Tadehagi auriculatum (DC.) H.Ohashi. Burma. Sylhet ，Chattuc k. C. B. Clarke 42702B (CAL ，cited in Ohashi 1973). October October 2003 Joumal of Japanese Botany Vo l. 78 No. 5 283

Reinwardtia Reinwardtia 6: 263 (1962); Sanjappa ， Leg. long). Ovary densely puberulent along both Ind. Ind. 164 (1 992). margins ， minutely stipitate. Pteroloma Pteroloma auriculatum (DC.) Miq. ， F l. Loments (5-) 7-8 サointed ， corky in Ind. Ind. Ba t. 1(1): 258 (1 855) ，ut “ Pteroloma peric 紅 p，glabrescent ，reticulate on lateral auriculatum auriculatum Desv."; Schind l. in Repert. surfaces ， with a3 -4 mm long stipe; articles Spec. Spec. Nov. Regni Veg. 21: 272 (1925) ， ut almost quadrangular ，5-7 mm long ，5-7 mm Pteroloma Pteroloma auriculata (DC.) Schind l. & in wide. Repert. Repert. Spec. Nov. Regni Veg. Beih. 49: 37 Dist r.: Mauritius ，Seychelles ，Mascarene ， (1928). (1928). India (Bengal ，Andaman Islands) ，Bangla- Pteroloma Pteroloma tl 匂uetrum (L.) Desv. ex Benth. desh ，Myanmar. subsp. subsp. auriculatum (DC.) H.Ohashi in J. Jpn. Specimens examined: Mauritius. Sieber (L); Bo t. 41: 95 (1 966). Bourbon 576 (L). Seyche Il es. E. P. Wright s. n. (K). Tadehagi Tadehagi triquetrum subsp. auriculatum India. Roxburgh s. n. (K); Benga l. 23 Feb.1872. C. B. Clarke Clarke 16905 (K); Great Cocos Island. Prain in 1889 (DC.) (DC.) H.Ohashi in Ginkgoana 1: 293 ，t. 41b (L); (L); Moolmyne. H .I 5688M (K); S. Andaman. Heinnig (1973); (1973); Polhill in Fl. Mascareignes ， 80. Leg.: 590 (BO). Bangladesh. Sylhet ，Chattuck 200. 13 Jan. 107 ， p l. 36 (1 990). 1886. C. B. Clarke 42702A (K). Myanmar. Shrub. Shrub. Leaves 1-foliolate. Stipules n訂- Tenasserium. Helfer 1648 (GH ，K ， L); Pegu. Scott s. n. rowly ovate ， 1-1.5 cm long ，3 -4 mm wide ， (L). acute ， glabrous ， conspicuously striate. In the original description the loments of Petioles Petioles 1-3 cm long ， winged ， sp 紅 sely Desmodium auriculatum DC. were charac- appressed appressed pubescent; wings 1-4 mm wide at terized as “ leguminibus glabe 町 imis subcori- broadest broadest part ， almost glabrous ， joined to the aceis" and “Articuli leguminis 4- 5 maturi facile facile secedentes." Also ， the lea f1 et was stipel stipel at tip; stipels curved ，glabrous. Lea f1 et 旬 blade blade n紅 Towly ovate or ovate ，3-11 cm “foliolis ovatis petiolo quadruplo longiori- long ，2-4 cm wide ， acute at apex ， rounded to bus." The type specimen was noted as origi 回 slightly slightly cordate at base ， upper surface nating from Timor and may be 紅 the name glabrous ， lower surface sp 紅 sely appressed “ Pteroloma auriculatum Desv!" (Candolle pubescent pubescent on nerves ，sparsely appressed cili- 1825) or “ Heteroloma auriculatum Desv- ate ate along margin ，lateral nerves altemate ， aux" (Candolle 1826). The type is apparently 10-13 pairs ， looped within margin. different from the specimen of Desmodium Inflorescences Inflorescences pseudoracemose ， to ca. 30 auriculatum DC. in Candolle: Prodromi cm long ， densely f1 owered ，2 f1 0wers with Herbarium in G (IDC micro-edition ， sheet 1-2 1-2 f1 0wer buds per node ， rachis with uni- no. 381) ， because the specimen in G has formly formly patent minute hooked hairs. Primary pubescent . loments bearing 2-4 remaining bracts bracts ovate ，6-7 mm long ， ca. 3 mm wide ， articles and shorter leaflet-blades. The mi- striate ， puberulent; secondary bracts subu- crofiche specimen bears an label as “Hetero- late ， ca. 1.5 mm long. Pedicels 5-7 mm long ， loma αuriculatum. Timor. Museum de Paris covered covered densely with patent minute hooked 1821". I am not sure whether the label was hairs. hairs. Bracteoles subulate ， ca. 1 mm long. written by Desvaux ， Candolle or some other Calyx appressed pubescent ， with minute persons. hooked hairs ， 4-10bed; lateral lobes triangu- Schindler examined the original specimen lar ， ca. 2.5 mm long; the lowest lobe nar- on the microfiche and determined it as rowly triangular ， ca. 3 mm long. Standard Pteroloma tl 匂uetrum in 1925.1 examined a incurved ，wings 6-7 mm long ， longer than probable duplicate of Desmodium auricula- the the keel ， long clawed (ca. 2 mm long) ， keel tum of the Prodromi Herbarium in L (Fig. 8). distinctly distinctly auriculate ， long clawed (ca. 2 mm It is referable to Tadeh α gi triquetrum subsp. 284 植物研究雑誌第78 巻第5号 平成15 年10 月 trzquetrum. trzquetrum. Desmodium triquetrum (L.) DC. ，r. Prod 2: The type must be in P as cited above. 326 (1825); Wight & Am. ，Prodr. F1. Ind.

Schindler Schindler (1928) ，Meeuwen (1 962) and Orien t.: 224 (1834) ， p. p. ，exc 1. D. al ωat ， ωum η1; Polhill Polhill (1 990) recorded that the type is in P. Baker 泊inH蜘∞O 依k.ムO F日1. Bri 出t. Ind. 2: 224 (1 876) ， As pointed out by Meeuwen (1962) ， the type p. p. ，exc 1. D. alatum et D. auriculatum. must must be erroneously located. ]j α dehagi Pteroloma triquetrum (L.) Desv. ex Benth. auriculatum auriculatum has so far been not known in in Miq. ， P 1. Jungh.: 220 (1 852) ，p. ，exc 1. P. Timo r. α latum. Distr. Distr. (sp よ Sri Lanka ， India ， Nepal ， エ2-4. Tadehagi robustum Pedley in Bhutan ，Bangladesh ，Myanmar ，Thailand ， Austrobaileya Austrobaileya 5: 258 (1999) [Type: Westem Indochina ，Malesia ， S. & SW China ，Taiwan ， Australia. Australia. Mitchell Plateau (c. 1 km N of the Ryukyus (I s. Ishigaki ，introduced) ， N. CRA mining camp) ，14 0 49"S 125 0 40"E. 14 Australia(?) ， and Pacific (New Caledonia). Jan. Jan. 1982. B. R. Maslin 5085 (PERTH holo. Wight and Am ott (1834) made the cir- n. n. v.; K iso.)]. cumscription of Desmodium 〆 triquetrum (L.) Description Description see Pedley (1999). DC. broader than that of Candolle (1 825) by An isotype of this species in Kew does not inclusion of D. α latum DC. and D. have have mature loments ， but the loments 訂 e pseudotriquetrum DC. to its synonymy. shortly shortly stiped and sparsely pubescent on the Bentham (1852) followed the broader con-

lateral lateral surfaces. The leaflet 回 blades of T. cept under Pteroloma. Baker (1876) added robustum robustum were characterized as oblong or D. auriculatum DC. into D. triquetrum in the ovate ovate and 2-2.5 times longer than wide; the broader concept as a synonym. Kurz (1 876) ，

pods pods have 4- 5 articles and 紅 e sp 紅 sely pu 圃 on the other hand ，recognized the typical D. bescent bescent on the lateral surfaces; the articles in triquetrum as v紅 . genuinum and D. pseu- the the middle part of the loments 訂 e larger than dotriquetrum with D. alatum as D. trique- those those in T. triquetrum; i. e. ，ca. 5 mm long trum var. pseudotriquetrum. Further ， Prain and and 8 mm wide in T. robustum against 2- (1 897) revised these taxa as subspecies of 3.5 3.5 mm long and 4-6 mm wide in T. D. triquetrum and this concept for D. triquetrum triquetrum (Pedley 1999). tl 匂uetrum has been accepted by 乱1eeuwen Distr.: Distr.: Endemic to N. Australia. (1 962) ， Ohashi (1 966 as Pteroloma ， 1973 as Specimens Specimens examined: Australia. Western Australia. Tadehagi) ， Sanjappa (1 992) and others as Amax Crusher ，Mitchell Plateau ，N.W. Kimberley. K. cited for the co 町 esponding subspecies in the F. F. Kenneally 7027 (K); B. R. Maslin 5085 (K iso ふ synonymic lists in this paper ， although Schindler Schindler (1924) recognized them ， as in 2・2-5. Tadehagi triquetrum (L.) H. Candolle (1825) ， as distinct species under Ohashi Ohashi in Ginkgoana 1: 290 (1 973) ， p. p. ， Pteroloma. The previous concepts are ，how- exc 1. subspp.α latum ， auriculatum ， et ever ， revised in this paper as recognizing rodgeri; rodgeri; Dy Phon in Dy Phon & a1.， Fl. three independent species ， Tadehagi alatum ， Camb. Laos Vietn. 27: 50 (1 994) ，p. ，exc l. T. αuriculatum ， and T. tl 匂uetrum ， of which subsp. subsp. rodgeri; Yang & Huang in F1. Reipub. T. tl 匂uetrum includes two subspecies plus Popu 1. Sin. 41: 63 (1995). two hybrids. Hedysarum triquetrum L. ， Sp. P1.: 746 Tadehagi triquetrum is divided into two (1 753) [Type: Herb. Hermann 4: 18 ， No. subspecies by difference in mo 中hology ，dis- 286 ，left-hand (BM ，n. v.). Lectotypifi- tribution and habi t. It is distributed in Asia cation cation by Pedley in Turland and Jarvis from India ， Sri Lanka through SE. Asia ， (1 997)]. China to Taiwan and New Caledonia. The October October 2003 Joumal of Japanese Botany Vo l. 78 No. 5 285 distribution distribution pattem differs between subsp. hair density seems to be continous between triquetrum triquetrum and subsp. pseudotriquetrum; the pubescent to glabrous in New Guinea and former former is not found in the Himalayas and is the Philippines.

r紅 'e in Malesia ， whilst the latter is more re 回 The leaflet-blade of 14α dehagi triquetrum stricted stricted being absent from Sri Lanka ， shows a different tendency in shape between Bangladesh ，Myanmar ，Thailand ，Indochina ， subsp. triquetrum and subsp. pseudotrique- and Pacific (New Caledonia) ， while rare in trum (Ohashi 1973). The blades of subsp. Taiwan. Taiwan. Moreover ， the two subspecies differ triquetrum are usually three to six times in in habi t: subsp. t1 匂uetrum is usually erect longer than broad ， while those of subsp. but but subsp. pseudotriquetrum is usually pros- pseudotriquetrum less than three times trate trate to ascending. longer than broad. The blades of subsp. Tadehagi Tadehagi triquetrum in the present cir- triquetrum variable especially in the cumscription cumscription shows great morphological Philippines ，i. e. ， 13-18 cm long ，O.ι0.7cm variation variation in density of hairiness on loments wide (20-25 times longer than broad) in and shape of lea f1 et-blades in New Guinea ， Ramos & Edano Bur. Sci. 44688 (A (Fig. 9) ， the the Philippines and Timor. K ，NY ，US) ， 11-19.5 cm long ，1- 1. 5 cm Degree of hair density on the lateral sur- wide (11-13 times longer than broad) in face face of loments varies greatly especially in Nemoto et al. 10188 (TUS) ， or 13-13.5 cm New Guinea (with examples) and the Philip- long ， 1.2-1.5 cm wide (9-10 times longer pines pines from entirely glabrous (e. g. ， Reeder than broad) in Ramos & Edano Bur. Sc i. 865 ， Brass 24360 ， Henty NGF20600) ， 39222 (A ，K ，L ， US). A specimen from sparsely sparsely pubescent (e. g. ， Brass 27226 ， Timor ，Cinatti 113 (L) is an extremely large Leach & Galore NGF33266 ，Wiakabu & individual having lea f1 et-blades 20 cm long A.M.C. 73580) through moderately dense and 6.5 cm wide ，stipules to 2.5 cm long and pubescent pubescent (e. g. ， Brass 7965 ， Pullen 7245) to 1 cm wide ，and the inflorescence to 36 cm very very densely hirsute (e. g. ，Anta 49 ，Anta 56 ， long (in flower) ， although its f1 0wers and Brass Brass 7652 ，Gebo UPNG312 ，Henty & Katik bracts are normal in size. NGF38651 ， Pullen 6766 ， Saunders 47). The remarkable variation in Tadehagi It It has been regarded. as an important diag- triquetrum in N ew Guinea ， the Philippines nostic nostic character between subsp. t1 匂uetrum and Timor may be derived from evolutionary and subsp. pseudotriquetrum whether hairy events ， although there remain further prob- or or glabrous on the lateral surfaces of lems whether the variation correlates with loments. loments. Both are ，however ， always densely any conditions in their habitats. sericeous sericeous on the sutures of lomen t. Candolle Specimens of Tadehagi triquetrum from (1825) (1825) described the loment of Desmodium the Lesser Sunda Islands are rather uniform triquetrum triquetrum as “ leguminibus sessilibus hirsu- in having leaves similar to subsp. pseudo- tis" tis" and D. pseudotriquetrum as "legumini- triquetrum ， while their loments are pubes- bus bus glabris ad suturam utramque adpresse cent on the lateral surfaces as in subsp. villoso-ciliatis". villoso-ciliatis". The hairy loments develop t1 匂uetrum. An example for such specimens from the pubescent ov 紅 y， while glabrous is the so-called isotype of Desmodium loments loments from glabrous ov 紅 y. The thinly pu- auriculatum DC. from Timor (L) (Fig. 8). 1 bescent bescent pods 訂 e derived from pubescent pis- determined it Tadehagi t1 匂uetrum subsp. tils tils which become less hairy during develop- triquetrum in 1999. This form is similar to a men t. The loment character is ，therefore ， hybrid between subsp. t1 匂uetrum and subsp. useful useful for segregation two taxa within pseudotriquetrum that will be discussed Tadehagi Tadehagi triquetrum ， although the degree of later ， but 1 postpone decision on the hybrid- 286 286 植物研究雑誌第78 巻第5号 平成15 年 10 月

Fig. Fig. 8. Tadehagi triquetrum (L.) H.Ohashi subsp. triquetrum. a: So-called isotype of Desmodium auriculatum auriculatum DC. Timor. Unknown collector (L). b: Infructescence enlarged from fig. a.

ism ism for the specimens from the Lesser Sunda lateral surfaces; leaflet-blade usually more Islands Islands cited in this paper. Further study is than 3 times longer than broad ，linear to needed needed on the basis of ample materia l. narrowly oblong-ovate or narrowly ovate ...・ ...subsp. triquetrum Key to the subspecies of Tadehagi triquetrum: triquetrum: 2-2-5- 1. subsp. triquetrum: H.Ohashi in 1. 1. Loments glabrous on lateral surfaces; leaf- Ginkgoana 1: 296 ， p l. 42a ，figs. 81 ，82 ， 85 let-blade let-blade less than 3 times longer than (1 973); Dy Phon in Dy Phon & al.， Fl. broad ，usually ovate or narrowly ovate. Camb. Laos Vietn. 27: 50 ， pl. 11:4-5 (1 994).

...・ H ・..… subsp. pseudotriquetrum Hedysarum alatum Roxb. ，[Hort. Beng. 56 1. 1. Loments densely to sparsely pubescent on (1 814) ， nom. nud.] Fl. Ind. ed. 2， 3: 348 October October 2003 Joumal of Japanese Botany Vo l. 78 No. 5 287

(1 832) (白 de Prain 1897). Wiakabu & A. M.C. 73580 (A ，K ， L). Philippines. Desmodium αcrocarpum Hance in J. Bo t. Rarnos Luzon. & Edano Bu r. Sc i. (A 44688 ，K ，NY ， 7: 7: 473 (1 848) [Type: Ins. Hong-Kong US); Mindanao. Rarnos & Edano Bur. Sci. (A 39222 ， K，L ， US); Culion. Me ロill 519 (GH ，NY ， US); Chinensium. Chinensium. Hance? ， Herb. unknown]. Zarnbales Prov.. Masinloc Town. Coto. S. side of Sul Desmodium triquetrum (L.) DC. var. Lauis River. above Coto Mines. 15 0 35'N. 120 0 05'E. genuinum Kurz in 1. Asia t. Soc. Bengal ， P t. al t. 15-170 rn. T. Nernoto & al. 10188 (TUS). 2，Na t. His t. 45: 232 (1877). Records of distribution of this subspecies Desmodium tl 匂uetrum (L.) DC. subsp. from Australia (cf. Meeuwen 1962 ， Ohashi genuina genuina Prain in 1. Asia t. Soc. Bengal ， P t. 2， 1973) may be doubtful (Pedley 1999). Na t. His t. 66: 390 (1897). Pteroloma Pteroloma tl 匂uetrum (L.) Desv. ex Benth. 2-2-5-2. Tadehagi triquetrum (L.) H. s. s. S.: Schind l. in Repert. Sp. Nov. 20: 271 Ohashi subsp. pseudotriquetrum (DC.) (1924). (1924). H.Ohashi in Ginkgoana 1: 295 ， t. 42b Desmodium tl 匂uetrum (L.) DC. subsp. (1 973); Huang & H.Ohashi in Li & al.， Fl. triquetrum: triquetrum: Meeuwen in Reinwardtia 6: 262 Taiwan 3: 389 ，t. 650 (1 977); H.Ohashi & al. (1 962); Verdc. ，Man. New Guinea Leg.: 410 in Sc i. Rep. Tohoku Univ. ser. 4， 38: 329 (1 979); Sanjappa ， Leg. Ind. 164 (1 992). (1984) & 39: 239 (1 988); Huang & H.Ohashi Dist r. (subsp. triquetrum): Sri Lanka ， in Huang & al.， Fl. Taiwan ed. 2， 3: 369 ，t. India ， Bangladesh ， Myanmar ， Thailand ， 192 (1 993). Indochina ，Malesia ，S & SW China ，Taiwan Desmodium pseudotriquetrum DC. in (new record ，rare) ，Ryukyu (l s. Ishigaki ，in- Ann. Sci. Na t. Paris 4: 100 (1 825) [Type: troduced) ， N. Australia(?) ，and Pacific (New Nepa l. Wallich (G holo. ， n. V. ，A iso. p. p.)] Caledonia) Caledonia) . & Mem. Leg.: 320 (1 826); Me 町. in Philipp. Specirnens Specirnens exarnined frorn Japan ，Taiwan ， and J. Sci. ， C. 5: 87 (1 910) & in Trans. Ame r. Malesia Malesia (exc l. other regions): Japan. Ryukyu ，Is. Phi l. Soc. 24: 201 (1 935). Ishigaki ，Hirakubo. In a stock farrn. 8 May 1954. T. Desmodium triquetrum (L.) DC. var. Arnano Arnano 7352 (RYU). Taiwan (new record). Il an: pseudo t1 匂uetrum (DC.) Kurz in J. Asia t. Yuan-shan ，2.5 krn SW of I-lan City ，24 0 45'-121 0 43 ヘ srnall A srnall hill of about 40 rn high. C. C. Hsu (TAI 8382 ， Soc. Bengal ， P t. 2，Na t. His t. 45: 232 (1877). TUS). TUS). Lesser Sunda Is. Fl ores. Kooy 117b (L); Desmodium tl 匂uetrum subsp. pseudo- Schrnutz Schrnutz 237 (L); Schouten 99 (L); Verhe 討en 398 (L) ， triquetrum (DC.) Prain in J. Asia t. Soc. Verheijen Verheijen 399 (L) ， Verheijen 4417 (L); Tirnor. Bengal ， P t. 2，Na t. His t. 66: 390 (1 897); Unknown Unknown collector (L ， G-DC as Desmodium Meeuwen in Reinwardtia 6: 263 (1 962); auriculatum auriculatum DC. in Candolle: Prodrorni Herbariurn ， Verdc. ，Man. New Guinea Leg.: 410 (1 979); rnicro-edition IDC rnicro-edition sheet no. 381) ，Bloernbergen 3366 (K ，L) ， Walsh 17 (A ，BO) ，Cinatti 113 (L) ，Friedberg Grierson & Long ， F l. Bhutan 1(3): 669 195 195 (L) ，Wiriadinata 2969 (L); Steenis Bessilau. 18593 (1987); Sa 吋appa ， Leg. Ind. 164 (1992). (L); (L); Wet 訂. Steenis 3610 (K) ，Elbert 4372 (L) ，Elbert Pteroloma pseudotl 匂uetrum (DC.) 4483 4483 (L). New Guinea. Road to biv. Mok. fores t. Anta Schind l. in Repert. Sp. Nov. 20: 272 (1924). 49 49 (L); Road to Kp. Kakajan. Anta 56 (K ，L); Papua. Tadehagi Tadehagi pseudotriquetrum (DC.) H. Milne Milne Bay Distric t. Brass 24360 (A ，K ，L ，US); Lake Daviurnbu ， Middle Fl y River. Brass 7652 (A ，L); Ohashi in 1. Jpn. Bo t. 70: 116 (1 995); Yang Middle Middle Fl y Ri ver. Brass 7965 (A ，K ，L); Hornbrurn & Huang in Fl. Reipub. Popu l. Sin. 41: 65 Bluff. Bluff. Gebo UPNG 312 (L); Aru fi. Henty & Katik (1 995). NGF38651 NGF38651 (L); Tufi ， al t. 120 rn. Leach & Galore Distr. (subsp. pseudotriquetrum): India ， NGF33266 NGF33266 (L); Goragatabu Creek area. Pu llen 6766 Nepal ，Bhutan ， Malesia (r 紅 e in New Guinea (A ，K ，L); ca. .3 rniles S. of Morehead Patrol Pos t. Pullen Pullen 7245 (A ，K ，L); ca. 0.5 rnile frorn Kasiawa on and Philippines) ， S & SW China ， and Bekalana Bekalana trac k. Tufi Subdistric t. al t. ca. 200 ft. Taiwan. Saunders Saunders 47 (K ， L); Beside the Haus Tarnbaran. Specirnens exarnined (addition to Ohashi 1973): 288 288 植物研究雑誌第78 巻第5号 平成15 年10 月

Fig. Fig. 9. Tadehagi triquetrum (L.) H.Ohashi subsp. triquetrum (D C.) H.Ohashi. a: Philippines. Luzon. Ramos & Edano Bur. Sci. 44688 (A). b: Young infructescence enlarged from fig. a， showing densely pilose loments. loments.

India. India. Assam. Chand 3604 (L) ，Chand 8253 (L) ， Koelz The typical form of subsp. pseudotrique- 24228 24228 (L) ， Koelz 28746 (L) ， Masters s. n. (L); Benga l. trum (the type from Nepal) has ovate to nar- Biswas Biswas 6480 (A); Darjeeling. Cave 11 (GH); Kumaon. rowly ovate leaflet-blades. It is common in Strachey Strachey & Winterbottom s. n. (GH); Manipur. Bullock Bullock 637 (K); Mussoorie. Stewart 16120 (A). the Himalayas ， China and Taiwan ， but rare Myanmar. Rangoon. Dickason 5087 (A); Namti River in New Guinea and the Philippines. One Valley ，Vicinity of Myitkyina. R. O. Belsher 623 (K); specimen ， Williams' collector 1414 (GH ， Camp Landies area ， Myitkyina region. R.O. Belsher NY) ，is the typical form from the Philippines 730 (K). New Guinea. Dobodura-Buna 町 ea. Reeder (Fig. 10). 865 865 (A ， US); Milne Bay Distric t. Brass 24360 (K ，L ， US); US); Onasinkian. Henty NGF20600 (L). Philippines. Luzon. Luzon. Lepanto. Merrill 4477 (K ，NY ， US); Luzon. Williams Williams collector 1414 (GH ，K ，NY ， US). October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 289

Fig. Fig. 10. Tadehagi triquetrum (L.) H.Ohashi subsp. pseudotriquetrum (DC.) H.Ohashi. a: Philippines. Luzon. Luzon. Williams collector 1414 (GH). b: Infructescence enlarged from fig. a，showing glabrous loments on lateral surfaces.

2-2-5-3. 2-2-5-3. Presumable intraspecific .hybrids the pods are referable to T. triquetrum subsp. in in Tadehagi tl 匂uetrum. pseudotl 匂uetrum because they 紅 e glabrous Intraspecific Intraspecific hybrids are recorded only in on the lateral surfaces and appressed hairy Desmodium and Tadehagi in the subtribe on the m 紅 gins (Fig. 11). A similar speci- Desmodiinae. Desmodiinae. Schindler (1924) recognized a men ， also from the Philippines ，Ramos & hybrid hybrid in Tadehagi (as Pteroloma) for the Edano Bur. Sci. 29494 (K) ，was determined first first time as “ Pteroloma pseudotl 匂uetrum x by Schindler on 28 Feb. 1926 as “Pterolom α ?triquetrum" ?triquetrum" based on a specimen from the pseudotriquetrum x ?α latum or ?triquetrum" Philippines ，Ramos Bur. Sci. 1036. He con- with the note “ legumina sterilia. Pt. alatum sidered sidered that the loments and calyx of the in insulis Philippine non reperitier" (Fig. 12). specimen 訂 e similar to P. pseudotriquetrum ， However ， the loments on this specimen 訂 e but but the leaves 紅 e similar to P. triquetrum immature and 1 cannot be certain whether (erroneously (erroneously as P. alatum). 1 examined a du- they would have produced seeds or no t. The plicate plicate of this specimen in GH in which the shape of leaflet-blade is similar to typical leaves leaves of the specimen 征 'e similar to subsp. tf 匂uetrum ，i. e. ， narrowly oblong or Tadehagi Tadehagi triquetrum subsp. triquetrum ， but narrowly ovate (9-10 cm long ， 1.2-2 cm 290 290 植物研究雑誌第78 巻第5号 平成15 年10 月

Fig. Fig. 11. Tadehagi triquetrum (L.) H.Ohashi nothosubsp. philippinensis H.Ohashi. a: Philippines. Luzon. Prov. Prov. Rizal ， Bosoboso. July 1906. M. Ramos Bur. Sci. no. 1036 (GH-holotype). A duplicate of voucher specimen specimen of Pteroloma pseudotriquetrum x ?triquetrum determined by Schindler. b: Infructescence with loments loments glabrous on lateral surfaces ， enlarged from fig ，a.

wide) ， and the loments 訂 e glabrous on lat- triquetrum (Fig. 13). The specimen ， Brass eral eral surfaces. 27226 (K) from New Guinea ， has ovate leaf- The following specimens are also interme- let-blades (2.2 -4 .7 cm long ， 1. 1- 1. 8 cm diate diate between subsp. triquetrum and subsp. wide; less than 3 times longer than broad) ， pseudotriquetrum: pseudotriquetrum: Ramos Bur. Sci. 22685 which is similar to those of subsp. (GH) has filiform leaves (13 cm long ，1. 5 cm pseudotriquetrum ， while its pods is sparsely wide; wide; ca. 8 times longer than broad) and pubescent on the lateral surfaces. loments loments glabrous on the lateral surfaces; 1 regard these intermediate forms as hy- Vidal Vidal 2610 (A) has also leaflets similar to brids between Tadehagi triquetrum subsp. subsp. subsp. triquetrum ， but pods are similar to triquetrum and subsp. pseudotriquetrum ， as subsp. subsp. pseudotriquetrum ， and Edano PNH they occurs only in the Philippines and New 14138 14138 (A ， L) has loments as subsp. pseudo- Guinea where both subspecies are known. triquetrum ， but leaflet-blade as subsp. Some problems regarding possible hybrids October October 2003 Journal of Japanese Botany Vo l. 78 No. 5 291

Fig. Fig. 12. Tadehagi triquetrum (L.) H.Ohashi nothosubsp. phil 争']J inensis H.Ohashi. a: Philippines. Luzon. Ri zal. Ramos & Edano Bur. Sci.29494 (K). A voucher specimen determined by Schindler as “ Pteroloma pseudotr 匂uetrum x?α latum or ?triquetrum~'. b: Y oung infructescence with loments glabrous on lateral surfaces ，enl 訂 ged from fig. a. remain remain in specimens from the Lesser Sunda Tadehagi triquetrum inter subsp. trique- Islands Islands in this species. trum et subsp. pseudotriquetrum quasi inter- media media et verisimiliter ex hybridatione harum triquetrum triquetrum (L.) H.Ohashi Tadeh~gi specierum orta ， ab ambobus fructibus nothosubsp. nothosubsp. philippinensis H.Ohashi ， hybr. abortivis abortivis differt ， quoad formam foliorum nov. nov. (Fig. 12) cum subsp~ triquetro sat congruens sed Pteroloma Pteroloma pseudotriquetrum x ?trique- leguminibus leguminibus lateralibus glabris. trum: trum: Schind l. in Repert. Sp. Nov. 20: 273 Specimens Specimens examined: Philippines. Luzon. Prov. (1 924). Rizal ， Bosoboso. July 1906. Ramos Bur. Sci. no. 1036 Tadehagi Tadehagi triquetrum su b， sp. triquetrum x (GH-holotype ，NY ， US); Prov. Ri l. za Ramos Bur. Sc i. subsp. subsp. pseudotriquetrum (DC.) H.Ohash i. 22685 (GH); Riza l. Ramos & Edano Bur. Sci. 29494 292 292 植物研究雑誌第78 巻第5号 平成15 年 10 月

(K ， US); Manila. Vidal 2610 (A ， K). lar to P. pseudotriquetrum ， but the leaves are The type ，Ramos Bur. Sci. 1036 (GH) ，was similar to P. triquetrum (erroneously as P. determined determined by Schindler (1924) as “Ptero- αlatum). Also ，1 confirmed that the leaves of loma pseudotriquetrum x ?triquetrum". He the specimen 紅 'e similar to Tadehagi considered considered that the pods and calyx are simi- tl 匂uetrum subsp. tl 匂uetrum ， but the pods

Fig. Fig. 13. Tadehagi triquetrum nothosubsp. philippinensis var. palawanensis H.Ohashi. a: Philippines. Philippines. Palawan: Edano ，4 Mar. 195 1. PNH14138 (A-holotype). b: A loment thinly pubescent pubescent on lateral surfaces ， enlarged from fig. a. October October 2003 Joumal of Japanese Botany Vo 1. 78 No. 5 293 are are referable to T. triquetrum subsp. pseudo- References t1 匂uetrum because they are glabrous on the Bailey C. D. ， Doyle J. J. ，K 吋ita T. ，Nemoto T. and lateral lateral surfaces and appressed hairy on the Ohashi H. 1997. The chloroplast rpl2 intron and margins. margins. Another specimen ， Vidal 2610 (A) ， ORF184 as phylogenetic markers in the legume tribe tribe Desmodieae. Sys t. Bo t. 22: 133-138. also also has leaves similar to triquetrum ， but Baker J. G. 1876. Desmodium. In: Hooker J. D.. The pods 紅 e similar to pseudo t1 匂uetrum. Fl ora of British 1ndia 2: 161-175. Ken t. Two forms are recognized in the hybrid: Balakrishnan N. P. 組 d Nair N. G. 1982. New taxa and i. i. Loments densely to sparsely pubescent on record from Saddle Peak ，Andaman 1slands. Bull. lateral lateral surfaces (as subsp. tl 匂uetrum) ， Bo t. 1ndia Surv. 24: 28-36. Bentham G. 1852. Leguminosae. In: Miquel F. A. W. ， leaflet-blade leaflet-blade less than 3 times longer than Plantae Junghuhnianae. pp. 205-269. Sythoff ， broad ，usually ov 御 or narrowly ovate (as Leiden. subsp. subsp. pseudotriquetrum) …・・・ . Candolle A. P. de 1825. CXLII 1. Desmodium D C. Prodromus Prodromus Systematis N aturalis Regni Vegetabilis ・H ・...var. phil 伊'P inensis i. i. Loments glabrous on lateral surfaces (as 2: 325-339. Treuttel et Wurtz ，Paris. subsp. subsp. pseudotriquetrum); leaflet-blade 一一一 1826. Memoires sur la famille des Legumine- uses. uses. Revue de la tribu des Heysarees. pp. 295- usually usually more than 3 times longer than 355. A. Belin ，Paris. broad ，linear to narrowly oblong-ovate or Dy Phon P. 1994. Tadehagi. In: Morat ， Ph. (ed よFlora

narrowly narrowly ovate (as subsp. triquetrum)... ・H ・ du Cambodge du Laos et du Vietnam 27. Dy Phon … var. palawanensis P. ， Ohashi H. and Vidal J. E.: Leguminosae ()- Papilionoideae 心 esmodieae. p. 47-54. Museum National d'Histoire Naturelle ，Paris. 2-2-5-i: 2-2-5-i: Tadehagi triquetrum nothosubsp. Ferguson Ferguson 1. K. and Skvarla J. 198 1. The pollen mor- philij フ'P inensis H.Ohashi var. philippinensis. phology of the subfamily Papilionoideae 2-2-5-ii: 2-2-5-ii: Tadehagi tl 匂uetrum nothosubsp. (Leguminosae). In: Polhill R. M. and Raven P.H. phil 伊'P inensis H.Ohashi var. palawanensis (eds よAdvances in Legume Systematics. pp. 859- 896. 896. Royal Botanic Gardens. Kew. H.Ohashi ，v 訂. nov. (Fig. 13) Hybrida Hybrida e Tadehage triquetro subsp. Goldblatt P. and Johnson D. E. (eds ふ 199 1. 1ndex to plant plant chromosome numbers 1988-89. Monographs triquetro triquetro et subsp. pseudotriquetro exorta ， in Systematic Botany from the Missouri Botanical sed sed varietas haec ab var. phil 伊vinensi differt Garden vo l. 40. foliis foliis subsp. pseudotriquetri et leguminibus 一一一 and 一一一 (eds.). 1998. 1ndex to plant chromo- subsp. subsp. triquetri. some numbers 199 4- 95. Monographs in Syste- Specimens Specimens examined: Philippines. Palawan: Edano matic Botany from the Missouri Botanical Garden vo l. 68. 4 Mar. 195 1. PNHI4138 (A-holotype ， L). New Kurz Kurz S. 1976. Contributions towards a knowledge of Guinea. Guinea. Fergusson 1s. Brass 27226 (A ，K ，L ， US). the the Burmese flora (continued). J. Asia t. Soc. Bengal ， P t. 2，Na t. His t. 45: 20 4- 310. 1 thank Dr. Bin Ye of the Tohoku Meeuwen M. S. Knaap-van 1962. Preliminary revi- University University for providing unpublished results sions of some genera of Malaysian Papilionaceae of of pollen morphology and Dr. Tadashi K 司ita V -A census of the genus Desmodium. Reinwardtia of of The University of Tokyo also for provid- 6: 239-276. ing ing unpublished molecular data. 1 wish to ex- Ohashi H. 1966. Nomenclatural notes on some taxa of Leguminosae. Leguminosae. J. Jpn. Bo t. 41: 95-96. press press my sincere thanks to the curators of A 一一一 1971. A taxonomic study of the tribe Coronilleae & GH ，BKF ，BO ，CAL ，K ，KYO ，L ，LAE ， (Leguminosae) ， with a special reference to pollen NY ，PNH ，TI ， and US for allowing me to morphology. 1. Fac. Sci. ， Univ. Tokyo ，Sec t. 1II work in their herbaria an d/ or for loaning to (Botany) 11: 25-92. me their specimens. Also ，1 thank Mr. 一一一 1973. The Asiatic Species of Desmodium and It s Allied Allied Genera (Leguminosae). 318 pp. ，76 pls. Gregory Gregory Kenicer of The University of Tokyo Ginkgoana Ginkgoana 1. Academia Scientific Book ，lnc. ， for for his reading the manuscrip t. Tokyo. 294 植物研究雑誌第78 巻第5号 平成15 年10 月

一一一 1982. The taxonomic position of Tadehagi Mahendra Pal Singh ，Dehra Dunn. rodgeri rodgeri (Legurninosae-Desrnodieae). J. Jpn. Bot. Schindler A. K. 1924. Ub er einige kleine Gattungen 57: 57: 26 4- 268. aus der Verwandschaft von Desmodium Desv. 一一一 1992. A new species of Desmodium Repert. Spec. Nov. Regni Veg. 20: 266-286. (Legurninosae) (Legurninosae) frorn Laos. J. Jpn. Bo t. 67: 320- 一一 1928. Die Desrnodiinen in der botanischen 323. 323. Li teratur nach Linne. Repert. Spec. Nov. Regni 一一一 1995. An enurneration of Chinese Desmodium Veg. Beih. 49: 1-37 1. and its allied genera (Leguminosae). 1. Jpn. Bo t. Schubert B. G. 1964. Anton K. Schindler. Taxon 13: 70: 70: 111-117. 7-10. 一一一， Polhill R. M. and Schubert B. G. 198 1. Tribe 9. Turland N. J. and Jarvis C. E. 1997. Typification of Desrnodieae Desrnodieae (Benth.) Hutch. (1 964). In: Polhill R. sorne Linnaean specific and varietal narnes in the M. and Raven P. H. (eds.) ，Advances in Legurne Legurninosae (Fabaceae). Taxon 46: 457 -4 85. Systernatics. Systernatics. 1. 292-300. Royal Botanic Gardens ， Wight R. and Arn ott G. A. W. 1834. Desmodium. Kew. Desv.; DC. In: Prodrornus Florae Peninsula Indiae Pedley Pedley L. 1999. Desmodium Desv. (Fabaceae) and re- Orientalis 1: 223-230. Parbury ，Allen ，& Co. ， lated lated genera in Australia: a taxonornic revision. London. Austrobaileya Austrobaileya 5(2): 209-261. Yang Y. C. and Huang P. H. 1995. Desrnodieae. In: Polhill Polhill R. M. 1990. 80. Leurnineuses. Flore des Lee S. K. (ed よ Flora Reipublicae Popularis Mascareigenes. Mascareigenes. The Sugar Industry Research Sinicae 41. Legurninosae (3) Papilionoideae 2. Institute ， Mauritius & a l. Science Press ，Beijing. Prain Prain D. 1897. Noviciae Indicae XV. Sorne additional Ye B. and Ohashi H. 2002. Morphology and polyrnor- Legurninosae. Legurninosae. J. Asia t. Soc. Bengal ， P t. 2，Na t. phisrn in the pollen of Christia and Uraria His t. 66: 347-518. (Legurninosae: Papilionoideae). J. Jpn. Bo t. 77: Sanjappa Sanjappa M. 1992. Legurnes of India. Bishen Singh 150-162.

大橋広好:マメ科タデハギ属の再検討と新属 Akschindlium の設立 1973 年にアジアのヌスピトハギ属とその近縁属 H.Ohashi およびタデハギ T. tr 匂uetrum (L.) を分類し ，Pteroloma 属に Droogmansia 属の 1 種 H.Ohashi の 6 種を独立種と認めた.これまで多く

Droogmansia god ψ， oyana (Kuntze) Schind l.を加え の地域のタデハギを調べてきたが，この種はフィ て独立属とし，タデハギ属 Tadehagi H.Ohashi と リピン，小スンダ列島およびニューギニアにおい 命名した.その後も新たな資料を加えてタデハギ て小葉の形と果実の毛の密度で特に著しい変異を 属について検討を続けた結果，本論文では 示すことが明らかになった.しかし，果実側面が Droogmansia god ザシ， oyana をAkschindlium H.Ohashi 無毛の subsp. pseudotriquetrum (D C.) H.Ohashi は として独立属とするとともに，タデハギ属を再構 葉身の形，生育形および分布のパターンがタデハ 成した . Akschindlium はタイとインドシナに分布 ギと異なるので，亜種として区別した.また，フィ する 1 種よりなる.この属名はヌスピトハギ連の リピンから記録されていた雑種 Pteroloma 分類に大きな貢献をしたドイツ人 Anton K. pseudotriquetrum x ?triquetrum: Schind l.をタデハギ Schindler 博士 (1879-1964) を記念した. タデハ と subsp. pseudotJ 匂uetrum の雑種と認め， ギ属 ，Akschindlium および Droogm αnsia の区別点 nothosubsp. philippinensis H.Ohashi と命名し， var. は本文に示した.タデハギ属としては Tadehagi palawanensis H.Ohashi を区別した.なお台湾にも alatum alatum (DC.) H.Ohashi ， T. andamanicum (Bal 北r. & 亜種タデハギがあることを記録した. N.G.Nair) N.G.Nair) S.P.Mathew ， T. auriculatum (DC.) (東北大学大学院理学研究科 H.Ohashi ， T. robustum Pedley ， T. rodgeri (Schind l.) 附属植物園津田記念館)