A Taxonomic Revision of Amphicarpaea (Leguminosae) Including a Pollen Morphological Comparison with Shuteria

Home , Amphicarpaea, Amphicarpaea bracteata, Dumasia, Glycine (plant), Phaseoleae, Pueraria

J. Jpn. Bot. 91 Suppl.: 231–249 (2016)

A taxonomic revision of Amphicarpaea (Leguminosae) including a pollen morphological comparison with Shuteria

a, b Hiroyoshi Ohashi * and Kazuaki Ohashi

aHerbarium TUS, Botanical Garden, Tohoku University, Sendai, 980-0862 JAPAN; bSchool of Pharmacy, Iwate Medical University, Yahaba, Iwate, 028-3694 JAPAN *Corresponding author: [email protected]

(Accepted on January 9, 2016)

Amphicarpaea is newly circumscribed based on the structure of the pollen apertures having linear grooves or slight colporus-like furrows instead of distinct colpori. Three species are recognized as the members of the genus: A. edgeworthii and A. ferruginea in Asia and A. bracteata in North America. Amphicarpaea africana having distinct tricolporate pollen grains is returned to Shuteria, and Amphicarpaea lineata in China is regarded as identical to Shuteria hirsuta. Amphicarpaea ferruginea Benth. is lectotypified.

Key words: Amphicarpaea, colporus-like furrows, Fabaceae, Phaseoleae, pollen aperture, pollen colporus, pollen morphology, Shuteria, Shuteria africana, typification.

Amphicarpaea is a papilionaceous genus flowers. belonging to the tribe Phaseoleae. It was Amphicarpaea was first placed in its named on the basis of the nature of A. bracteata taxonomic position between Apios and having different two kinds of fruits: aerial and Glycine (Nuttall 1818). It has been classified in subterranean fruits. In fact, the species has two Phaseoleae subtribe Glycininae (Bentham 1865, kinds of aerial pods and subterranean ones. Lackey 1981, Schrire 2005) or tribe Glycineae Amphicarpaea edgeworthii has the same habitat. (Hutchinson 1964). Evolutionary relationships The aerial pods are produced from the aerial between Amphicarpaea and other genera chasmogamous flowers and cleistogamous ones. among Glycininae have been problematic. The subterranean pods arise from cleistogamous Bentham (1865) classified Amphicarpaea flowers borne on the subterranean shoots between Cologania and Dumasia. Taubert elongating from the cotyledon axil and geocarpic (1894) united Amphicarpaea with Cologania. pods from cleistogamous flowers on terrestrial Hutchinson (1964) put the genus next to creeping shoots grown from the aerial axil of the Shuteria. Turner and Fearing (1965) supposed lower leaves. These two species, A. bracteata a probable evolutionary relationship between and A. edgeworthii, are annual. In contrast, A. Amphicarpaea and Shuteria. In fact, several ferruginea and A. africana are perennial, having species were transferred between the two two kinds of aerial pods produced from aerial genera, e.g., A. africana or S. africana, A. chasmogamous flowers or cleistogamous ones, ferruginea or S. ferruginea, or A. trisperma or but lacking subterranean or geocarpic pods. S. trisperma. Lackey (1981) considered that Amphicarpaea lineata has no cleistogamous Amphicarpaea is particularly closely allied to

—231— 232 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Shuteria, Dumasia and Cologania in its flowers. Pollen grains of Shuteria were treated by However, Lee and Hymowitz (2001) analyzed Thuan (1972) who excluded Shuteria anomala molecular phylogenetic relations in the subtribe Pamp. and S. longipes Franch. from this genus and showed that “Amphicarpaea was sister to based partly on pollen morphological characters. the clade comprising Glycine and Teramnus He transferred S. anomala to a synonym rather than to Cologania.” Cologania, Dumasia of Amphicarpaea edgeworthii, but did not and Shuteria showed remote relationships accommodate S. longipes to any genus. Now S. with Amphicarpaea in their analysis. In recent longipes is placed in Hylodesmum of the tribe molecular analyses of the phaseoloid groups Desmodieae, H. longipes (Franch.) H. Ohashi (Phaseoleae s.l.), Amphicarpaea belongs to & R. R. Mill (Ohashi and Mill 2000). Ferguson the “Core-Phaseoleae” clade, while Shuteria and Skvarla (1981) introduced the Shuteria is distant from the clade and grouped with a pollen grains as subprolate, margocolporate, and Kennediinae-Desmodieae clade (Schrire 2005, reticulate with granules in the lumina. Stefanović et al. 2009). This paper aims to revise Amphicarpaea, as Materials and Methods the genus needs to have a clear circumscription. Pollen materials were obtained from A comparative pollen morphological study herbarium specimens kept in E and TUS. is emphasized between Amphicarpaea The pollen grains for SEM observation were and Shuteria, because these are hardly acetolysed following the standard method distinguishable from each other in external (Erdtman 1960) and dehydrated in an ethanol morphology, but no discussion has been made series. The air-dried samples were coated with of their pollen morphological relationships. osmium with a Filgen NL-OPC 60A osmium Detailed morphological descriptions for A. plasma coater, and examined with a Hitachi edgeworthii and A. ferruginea are provided. SU-8010 scanning electron microscope at Amphicarpaea lineata is newly placed as a the Center for Electron Microscopy and Bio- synonym of S. hirsuta in this paper. Imaging Research, Iwate Medical University. Observations by SEM are performed at 2.0 kV. 1. Pollen morphology of Amphicarpaea and The polar and equatorial lengths were measured Shuteria on at least 20 grains of each specimen. Pollen Pollen grains of Amphicarpaea were terminology in our study generally follows recorded by Schively as early as in 1897 for A. Punt et al. (1994), Hoen (1999) and Hesse et al. comosa (= A. bracteata). Differences in pollen (2009). morphology in Amphicarpaea were first shown by Meeson (1974) based on observations in light Pollen descriptions microscopy among the three species recognized Amphicarpaea africana Harms (Fig. 1) by Turner and Fearing (1965): 3-colporate in A. Pollen grains tricolporate; polar axis 25.3 africana, 4-porate in A. bracteata, and 3-porate ± 2.0 µm (range: 21.2–27.7 µm) (n = 20), in A. edgeworthii. He stated that “the pollen of equatorial diameter 23.9 ± 1.9 µm (21.0–27.2 Amphicarpaea edgeworthii and A. bracteata µm) (n = 28), P/E = 1.07 ± 0.08 (0.88–1.20), appear to be modifications of the basic form of spheroidal, equatorial view elliptic, polar view A. africana by loss of the colpi and reticulated triangular to circular. Colpori 0.60 times as sexine and an increase of retention in the number long as the polar axis, ca. 1.8–3.6 µm wide at of pores.” Ohashi et al. (2005) accepted this equator, narrowing to pointed ends; colporus view and regarded that the pollen of the genus membrane almost smooth, margines with has a basically colporate structure. continuous tectum; endoaperture ca. 0.12 times December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 233

Fig. 1. Pollen grain of Amphicarpaea africana. A. Mesocolpium in equatorial view. B. Aperture in equatorial view. C. Polar view. D. Sculpture of mesocolpium. Voucher specimen: Congo, Semliki R. forest. Kassner Expedition 1908, no. 3088 (E). Scale bar: A, B, C = 10 µm; D = 1 µm.

Fig. 2. Pollen grain of Amphicarpaea bracteata. A. Mesocolpium in equatorial view. B. Aperture in equatorial view. C. Polar view. D. Sculpture of mesocolpium. Voucher specimen: U.S.A. Vermont. Franklin Co., Fletcher, alt. ca. 240 m. 19 Aug. 1983. Boufford & Wood 23201 (TUS). Scale bar: A, B, C = 10 µm; D = 1 µm. 234 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue as long as the polar axis. Sexine semitectate; not. Ohashi et al. (2005) supposed the structure sculpture of mesocolpium reticulate, perforate as rudimentary colpori and regarded the pollen around margines, lumina 0.5–3 µm in diameter as colporate. The grooves or colporus-like at mesocolpium, with columellae elements. furrows are, however, different from the typical Voucher specimen: Congo. Semliki R. forest. colporus, because they seem to have no obvious Kassner Expedition 1908, no. 3088 (E). ectoaperture. The pollen grains of Amphicarpaea africana differ from those of the other two species in Amphicarpaea edgeworthii Benth. (Fig. 3) having distinct colpi. We compared the pollen Pollen grains triporate usually with slight with that of Shuteria hirsuta, because A. africana colporus-like furrows; polar axis 22.5 ± 1.2 had been regarded as a member of Shuteria, µm (range: 20.6–25.5 µm) (n = 25), equatorial and S. hirsuta appears to be most similar to diameter 19.8 ± 1.0 µm (18.1–21.9 µm) (n = 26), Amphicarpaea among Shuteria. Pollen grains P/E = 1.13 ± 0.06 (1.08–1.27), prolate spheroidal of S. hirsuta and S. vestita were reported by to subspheroidal, equatorial view elliptic, polar Thuan (1972). We confirmed that pollen grains view circular. Aperture ca. 0.18 times as long of these species are distinctly tricolporate. The as the polar axis, margines continuous with colporus in the two Shuteria species has the mesocolpium. Sexine semitectate; sculpture of same structure as that of A. africana, but not of mesocolpium granulate to rugulate with fine A. bracteata and A. edgeworthii. fossula. Voucher specimen: Japan. Miyagi Pref., Sendai-city, Aobayama. 22 Sep. 1985. Amphicarpaea bracteata (L.) Fernald (Fig. 2) Nemoto 2881 (TUS). Pollen grains tetraporate sometimes with The pollen grains of this species (also as linear grooves or slight colporus-like furrows; Amphicarpaea trisperma, A. edgeworthii var. polar axis 28.4 ± 1.0 µm (range: 26.4–30.3 japonica or A. bracteata subsp. edgeworthii) µm) (n = 20), equatorial diameter 25.0 ± 1.4 have been described as 3-colporate, 3-porate µm (22.2–27.4 µm) (n = 20), P/E = 1.14 ± 0.06 or 4–5-porate (Ikuse 1956, Huang 1968, 1972, (1.06–1.22), prolate spheroidal to subspheroidal, Meeson 1974, Liu 1991, Guo et al. 2003, Ohashi equatorial view elliptic, polar view circular et al. 2005). Such divergent descriptions were to polygonal. Colporus-like furrows (when produced due to occurrence of the colporus-like observed) continuous with mesocolpium, structure which are often indistinguishable from lacking margines, 0.44 times as long as the the tectum (Ohashi et al. 2005). Variation of the polar axis and ca. 1.9–3.2 µm wide at equator; colpori was illustrated in Ohashi et al. (2005; fig. aperture ca. 0.16 times as long as the polar axis. 156A–C). After reexamination, the colpori in Sexine semitectate; sculpture of mesocolpium the previous description should be corrected as rugulate with fine fossula. Voucher specimens: colporus-like furrows; hence the pollen of this U.S.A. Vermont. Franklin Co., Fletcher, alt. ca. species is triporate. 240 m. 19 Aug. 1983. Boufford & Wood 23201 (TUS). Amphicarpaea ferruginea Benth. (Fig. 4) The pollen of this species has been regarded Pollen grains triporate with slight linear as 4-porate (Meeson 1974) or 4(–5)-colporate grooves without margines; polar axis 36.2 ± 1.5 (Ohashi et al. 2005). The linear colporus-like µm (range: 33.8–38.4 µm) (n = 23), equatorial furrow was shown in Ohashi et al. (2005; fig. diameter 24.5 ± 1.2 µm (23.2–27.1 µm) (n = 24), 156D, E). The pollen types are differently P/E = 1.48 ± 0.08 (1.32–1.60), oblate to prolate, judged as to whether the structure of the linear equatorial view oblong to elliptic, polar view grooves or colporus-like furrows are colpori or triangular. Aperture ca. 0.10 times as long as the December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 235

Fig. 3. Pollen grain of Amphicarpaea edgeworthii. A. Mesocolpium in equatorial view. B. Aperture in equatorial view. C. Polar view. D. Sculpture of mesocolpium. Voucher specimen: Japan. Miyagi Pref., Sendai-city, Aobayama. 22 Sep. 1985. Nemoto 2881 (TUS). Scale bar: A, B, C = 10 µm; D = 1 µm.

Fig. 4. Pollen grain of Amphicarpaea ferruginea. A. Mesocolpium in equatorial view. B. A perture in equatorial view. C. Polar view. D. Sculpture of mesocolpium. Voucher specimen: Nepal. Khodang Danda 2500 m – Latsu 2400 m. 29 Aug. 1972. Kanai, Hara & Ohba 722043 (TUS). Scale bar: A, B, C = 10 µm; D = 1 µm. 236 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue polar axis. Sexine semitectate; mesocolpium semitectate; sculpture of mesocolpium reticulate, with incomplete reticulum, polar area rugulate, polar area psilate to perforate, lumina 0.5–3 µm lumina 1–3 µm in diameter at mesocolpium, in diameter at mesocolpium, with columellae with columellae elements. Voucher specimen: elements. Voucher specimen: Nepal. Arun Nepal. Khodang Danda 2500 m – Latsu 2400 Valley. 7 Nov. 1981. Tateishi 8455 (TUS). m. 29 Aug. 1972. Kanai, Hara & Ohba 722043 (TUS). 2. Taxonomic treatment The pollen morphology of Amphicarpaea Turner and Fearing (1965) recognized three ferruginea is recorded here for the first time. The species in Amphicarpaea “each of which is structure of the apertures is similar to that of A. confined to a separate continent: A. africana in bracteata and A. edgeworthii in having slight Africa, A. edgeworthii in Asia, and A. bracteata linear grooves or indistinct furrow, but differs in North America”. Shuteria ferruginea (Benth.) from those of S. hirsuta and S. vestita. Baker was resurrected to Amphicarpaea as the second Asian species (Ohashi 1975). Schrire Shuteria hirsuta Baker (Fig. 5) (2005) summarized the genus consisting of “4–5 Pollen grains tricolporate; polar axis 23.6 spp.; C and E USA, SE Canada to Mexico (1 ± 0.9 µm (minimum–maximum: 22.4–24.8 sp.); E. Asia (Russia, Japan, Korea, China) to µm) (n = 24), equatorial diameter 21.1 ± 0.6 Himalaya (2–3 spp.); montane tropical C and E µm (20.0–22.0 µm) (n = 27), P/E = 1.12 ± 0.04 Africa (1 sp.).” Amphicarpaea lineata Chun & (1.05–1.23), prolate spheroidal to subspheroidal, T. C. Chen was adopted as the third member of equatorial view elliptic, polar view triangular to the genus in the most recent Flora of China (Sa circular. Colpi 0.68 times as long as the polar and Gilbert 2010). axis, ca. 2.8–4.0 µm wide at equator, narrowing Amphicarpaea has been distinguished from to pointed ends; colpus membrane granulate, Shuteria by lacking septa between seeds in margines with perforate tectum; endoaperture the legumes (Hutchinson 1964, Thuan 1972) ca. 0.14 times as long as the polar axis. Sexine or lacking bracteoles (Sa et al. 2010). These semitectate; sculpture of mesocolpium reticulate, characters used in the key to the genera are, polar area perforate, lumina 0.5–2 µm in however, useless for distinctions between diameter at mesocolpium, with columellae the two genera. Shuteria hirsuta has legumes elements. Voucher specimen: Nepal. Koshi without septa between the seeds. Amphicarpaea Zone, Sankhuwasabha Distr. 29 Aug. 1998. ferruginea is rarely bracteolate. Generally, Noshiro et al. 9810179 (TUS). Amphicarpaea has 2–3(–4)-seeded legumes, while Shuteria is 4–10 seeded. No clear Shuteria vestita Wight & Arn. (Fig. 6) demarcations have been found between both Pollen grains tricolporate; polar axis 26.1 genera in external morphology. ± 1.0 µm (minimum–maximum: 24.4–28.1 The pollen structure of Amphicarpaea is µm) (n = 20), equatorial diameter 22.3 ± 0.9 characteristic in lacking clear colporus but having µm (20.6–24.0 µm) (n = 23), P/E = 1.18 ± 0.04 slight linear grooves or slight colporus-like (1.13–1.24), prolate spheroidal to subspheroidal, furrows of which the margines are continuous equatorial view elliptic, polar view triangular to with mesocolpium. In light of the pollen circular. Colpi 0.76 times as long as the polar morphology described above, we consider that A. axis, ca. 2.1–3.6 µm wide at equator, narrowing africana should be returned to Shuteria, because to pointed ends; colpus membrane granulate, it differs from those of other Amphicarpaea margines with perforate tectum; endoaperture species in having distinct colpori. Amphicarpaea ca. 0.15 times as long as the polar axis. Sexine is proposed here as consisting of three species: A. December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 237

Fig. 5. Pollen grain of Shuteria hirsuta. A. Mesocolpium in equatorial view. B. Aperture in equatorial view. C. Polar view. D. Sculpture of mesocolpium. Voucher specimen: Nepal. Koshi Zone, Sankhuwasabha Distr. 19 Aug. 1998. Noshiro et al. 9810179 (TUS). Scale bar: A, B, C = 10 µm; D = 1 µm.

Fig. 6. Pollen grain of Shuteria vestita. A. Mesocolpium in equatorial view. B. Aperture in equatorial view. C. Polar view. D. Sculpture of mesocolpium. Voucher specimen: Nepal. Arun Valley. 7 Nov. 1981. Tateishi 8455 (TUS). Scale bar: A, B, C = 10 µm; D = 1 µm. 238 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue bracteata, A. edgeworthii and A. ferruginea. enclosed in calyx, petals absent or rudimental in aerial ones, stamens 10, free, rudimental, Amphicarpaea Elliott ex Nutt., Gen. N. ovary 1, 1–3-ovuled, style recurved. Pods Amer. Pl. 2: 113 (1818), as Amphicarpa, nom. produced from aerial chasmogamous flowers et orth. cons. [see Rickett & Stafleu in Taxon or aerial cleistogamous ones dehiscent, falcate 8: 296 (1959), Nicolson & al. in Taxon 33: or narrowly oblong, compressed, not septate, 316 (1984)]; DC., Prodr. 2: 383 (1825); Benth. margined, (1–)3–4-seeded, pericarp thin, pale in Benth. & Hook. f., Gen. Pl. 1: 451, 529 brown. Subterranean pods produced from axil of (1865); Hutch., Gen. Fl. Pl. 1: 449 (1964), as aerial shoots arising from the first or lowermost Amphicarpa; Turner & Fearing in Southwest. axil of stem or subterranean shoot from axil of Nat. 9: 207 (1965); Verdc. in Fl. Trop. E. Africa, cotyledon indehiscent, subglobose, 1–2-seeded, Legum. (Part 4): 511 (1971), as Amphicarpa; pericarp pale pink, membranous, glabrous. Lackey in Adv. Leg. Syst. 1: 319 (1981), as Pollen grains tri- or tetraporate; aperture often Amphicarpa; Lee & Hymowitz in Am. J. Bot. with linear grooves or slight colporus-like 88: 2071 (2001); H. Ohashi, Nemoto & Onodera furrows, lacking colporus; membrane of the in J. Jpn. Bot. 80: 153 (2005); Schrire in Lewis furrows same with that of mesocolpium. & al., Leg. World: 420 (2005). Chromosome numbers: 2n = 20, 22, and 40. Type species: Amphicarpa monoica Elliott Somatic counts for Amphicarpaea bracteata ex Nutt., nom. illeg. (Glycine monoica L., 2n = 20 (Turner and Fearing 1965) and 40 nom. illeg., Glycine bracteata L., Amphicarpa (Löve and Löve 1982) were probably erroneous bracteata (L.) Fernald). (Kumar and Hymowitz 1989). A somatic count Phaseolus subgen. Amphicarpaea (Elliott ex of 2n = 20 was also reported by Yeh et al. (1983) Nutt.) Eaton & Wright, N. Amer. Bot., ed. 8: 353 for A. edgeworthii. Espert et al. (2008) reported (1840), as Amphicarpa. 2n = 40 in A. bracteata and A. edgeworthii based Procumbent or twining herbs, annual or on more recent works by Kress et al. (2005) and perennial. Cotyledons hypogeal. Roots nodular. an unpublished data. Nishikawa (2008) cited Leaves petiolate, pinnately 3-foliolate, stipulate, 2n = ca. 22 for A. edgeworthii based on Fukui stipelate, rachis distinct, lateral nerves of and Suda’s report (1975), but the count was leaflets reaching margin. Inflorescences with tentative, not valid as stated by Fukui and Suda chasmogamous flowers axillary, pseudoracemes, (1975). The somatic chromosome numbers of sometimes with 1–2-flowered fascicles or Amphicarpaea may be 2n = 22 (x = 11 as the racemes with aerial cleistogamous flowers. basic number) on the basis of its systematic Primary bracts early deciduous or sometimes position among the subtribe Glycininae, but persistent; secondary bracts absent, very rarely further examinations are needed. present. Pedicels recurved pubescent. Bracteoles Distribution: Asia (Nepal, Sikkim, Bhutan, absent, very rarely present. Chasmogamous NE. India, Myanmar, Thailand, Vietnam, China, flowers: calyx tubular, 4-lobed, adaxial lobe Taiwan, Korea, Japan, and Far East Russia) and completely united, entire at apex; corolla America (SE. Canada, central to E. USA and glabrous, longer than calyx, standard obovate, Mexico). apex rounded; wings and keel with long claw, connate at center of lamina; stamens 10, Key to the species diadelphous, vexillary (adaxial) stamen free; 1. Leaflets attenuate to cuspidate, usually disk present around stalk of pistil; pistil stipitate, apiculate, texture papery or thickly papery; ovary 2–4-ovuled, style filiform, stigma small, perennial herb without subterranean pods; capitate, papillate. Cleistogamous flowers species of Asia ...... A. ferruginea December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 239

1. Leaflets obtuse or acute to shortly attenuate, Pedicels 1–4(–5) mm long, reflexed pubescent. texture membranous or thinly papery; annual Bracteoles absent. Chasmogamous flowers: herbs with subterranean pods ...... 2 calyx 4–6 mm long, ca. 2 mm wide; adaxial 2. Chasmogamous flowers: lateral calyx-lobes lobes united, broadly triangular, acuminate, 2–3 mm long, apex acuminate to caudate; 2–2.5 mm long; lateral lobes triangular, 1.5–2 bracts broadly ovate to orbicular, 1.5–4 mm mm long, abaxial lobe broadly triangular, wide; species of Asia ...... A. edgeworthii acuminate, 2–2.5 mm long; corolla white and 2. Chasmogamous flowers: lateral calyx-lobes lavender-tipped or light lavender to purple, 1–2 mm long, apex acute to acuminate; bracts 1.2–1.5 cm long; standard longer than wings and broadly to very broadly obcordate, 3–5 mm keels, 6–7 mm wide, without auricles; wings 12– wide; species of America ...... A. bracteata 13.5 mm long, lamina 4.5–5 mm long, auricled at base; keels as long as wings, lamina 5–5.5 mm 1. Amphicarpaea bracteata (L.) Fernald long. Stamens 12–13.5 mm long. Pistils as long in Rhodora 35: 276 (1933), as Amphicarpa as stamens. Pods from chasmogamous flowers bracteata; Rickett & Stafleu in Taxon 8: 296 stipitate (stipe 3–5 mm long), oblong, straight or (1959), as Amphicarpaea bracteata; Turner & falcate, 1.5–4 cm × 8–10 mm, apex acuminate; Fearing in Southwest. Nat. 9: 215 (1965); Isely, lateral surfaces glabrous or strigose, reticulate Vasc. Fl. S.E. U.S., Leguminosae: 177 (1990); H. nerved; sutures villous. Seeds 2–4, ca. 5 × 4 mm. Ohashi, Nemoto & Onodera in J. Jpn. Bot. 80: Subterranean legume, ellipsoid or subglobose, 133, fig. 156 pollen (2005), as subsp. bracteata. 0.8–1.5 cm long. Seed 1, large. – Glycine bracteata L., Sp. Pl. 2: 754 (1753) Chromosome numbers: 2n = 22 (Kumar and [Type: “Virginia madidis, umbrosis.” Kalm. Hymowitz 1989). Herb. Linn. No. 901.3 (LINN, lectotypification Distribution: SE. Canada, C. and E. USA, by Reveal & al. in Huntia 7: 230, 1987; Jarvis, and Mexico. Order Chaos: 544, 2007)]. See the synonymy in Turner and Fearing (1965). 2. Amphicarpaea edgeworthii Benth. Annual climbing herb. Stems aerial and in Miq., Pl. Jungh.: 231 (1852) [Type: subterranean from cotyledon axils. Aerial Himalaya. alt. 5–6000ft., shady woods, M. stem sparsely to densely, ascending to patent P. Edgeworth 169 (K000883275, K000883276 pubescent. Stipules persistent, ovate to –holotype) (Fig. 7)]; Baker in Hook. f., Fl. narrowly ovate, 2–5 mm long. Leaflets entire, Brit. Ind. 2: 181 (1876); Turner & Fearing in membranous or nearly so, ovate or broadly ovate Southwest. Naturalist 9: 212 (1965); Thuan, Fl. to broadly rhombic-ovate, sparsely pubescent Camb., Laos & Vietn. 17: 56 (1979); Y. T. Wei on nerves, lateral nerves 4–5, base rounded to in S. K. Lee, Fl. Reip. Popul. Sin. 41: 257, t. broadly cuneate or nearly truncate, apex acute 62 (1995); Zhang & al. in Biochem. Syst. Eco. to acuminate, terminal one 1–10 cm long, 1–8 33: 1246 genetics (2005); R. Sa & Gilbert, Fl. cm wide. Inflorescences with chasmogamous China 10: 249 (2010). – Falcata edgeworthii flowers pseudoracemes or racemes, axillary, (Benth.) Kuntze, Revis. Gen. Pl. 1: 182 (1891). simple or branched, sometimes branched, – A. bracteata (L.) Fernald subsp. edgeworthii 1.5–5 cm long, 5–18-flowered; flowers 1–2 per (Benth.) H. Ohashi in H. Hara, Fl. E. Himal.: 137 node. Primary bracts broadly obcordate to very (1966); H. Ohashi in Enum. Fl. Pl. Nepal 2: 105 broadly so, lower often enfolding the pedicel, (1979); H. Ohashi in Wild Herb. 2: 211 (1982); striate, 2–4 mm long, 3–5 mm wide, abaxial Sanjappa, Leg. India: 82 (1992); H. Ohashi in K. surface sericeous, adaxial surface glabrous, apex Iwats. & al., Fl. Jap. IIb: 277 (2001); H. Ohashi, rounded to bilobed. Secondary bracts absent. Nemoto & Onodera in J. Jpn. Bot. 80: 133, fig. 240 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Fig. 7. Holotype of Amphicarpaea edgeworthii Benth. K000883275 (©The Board of Trustees of the RBG, Kew). December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 241

156 pollen (2005). s.n. 26 Aug. 1904 (TI); Aizu. J. Matsumura s. n. A. edgeworthii Benth. var. japonica Oliv. in Aug. 1879 (TI); Aizu? K. Nemoto (?, n.v.)]. J. Linn. Soc., Bot. 9: 164 (1867) [Type: Japan. A. edgeworthii f. alba M. H. Lee in Korean J. Nagasaki. Fl. blue & white, amongst shrubs. Pl. Taxon. 13(2): 109 (1983). Sep. 1862. R. Oldham 910. K000883278– Glycine monoica auct. non L.: Thunb., Fl. holotype]; Franch. & Sav., Enum. Pl. Jap. 1: 107 Jap.: 283 (1784). (1873); Forbes & Hemsl. in J. Linn. Soc. Bot. Glycine javanica auct. non L.: Thunb. in 23: 188 (1887); Matsum. in Bot. Mag. (Tokyo) Trans. Linn. Soc. 2: 340 (1794). 16: 84 (1902); Ohwi, Fl. Jap.: 694 (1953), & Fl. Annual herb. Underground stems growing Jap. ed. Engl.: 571 (1965), & Fl. Jap. ed. rev. from axil of cotyledons filiform, to 30 cm 809 (1965); Kitam. & Murata, Col. Illust. Herb. long, white, sparsely reflexed hairy, with long Pl. 2: 117 (1961). – Falcata japonica (Oliv.) internode, with subterranean cleistogamous Kom. in Trudy Imp. S.-Peterburgsk. Bot. Sada flowers at node and apex. Terrestrial stems to 2 22: 630 (1904). Komar. in Acta Hort. Petrop. 22: m long, slender, green, with reflexed white or 630 (1904), Fl. Mansh.: 630 (1904); Makino, yellowish brown hairs, often producing axillary Illust. Fl. Jap.: 402, f. 1204 (1940); Bobrov in Fl. branches near the ground, filiform, green, with URSS 13: 527 (1948). – F. comosa var. japonica subterranean cleistogamous flowers at apex. (Oliv.) Makino in Bot. Mag. (Tokyo) 20: 82 Stipules ovate to broadly ovate, 3–5 mm long, (1906); Makino & Nemoto, Nippon Shokubutsu abaxial surface pubescent at base, ciliate, with Sôran (Fl. Jap.), ed. 2, 569 (1931). – A. japonica 6–8 parallel nerves, apex acute. Petioles 1–5.5 (Oliv.) B. Fedtsch. in Trudy Imp. S.-Peterburgsk. cm long, reflexed pubescent; stipels narrowly Bot. Sada 31(1): 155 (1912); N. S. Pavlova in ovate, ca. 1 mm long, early deciduous, glabrous, Kharkev., Pl. Vasc. Or. Extr. Soviet. 4: 208, t. 16 ciliate. Terminal leaflet rhomboid-ovate or V (1989). – A. bracteata subsp. edgeworthii var. broadly ovate, (2–)3–9 × (1.5–)2–6 cm, base japonica (Oliver) H. Ohashi in H. Hara, Fl. E. rounded, broadly cuneate or nearly truncate, Himal.: 139 (1966); T. C. Huang & H. Ohashi, apex acute or obtuse, sometimes acuminate, both Fl. Taiwan 3: 170, t. 517 (1977), & Fl. Taiwan surfaces white villous, appressed ciliate; rachis ed. 2, 3: 198, t. 88 (1993). 0.5–1.5 cm long; lateral leaflets smaller than Shuteria trisperma Miq. in Ann. Mus. Bot. the terminal one, ovate or ovate-rhombic, base Lugd.-Bat. 3: 51 (1867) [Syntype: Japan. obliquely rounded. Siebold, Buerger, Mohnike; in vallibus montis Inflorescences with chasmogamous flowers Sata Toge prope Kifura Sawa ins. Nippon, pseudoraceme or raceme, axillary, 1–2 per node, Pierot. (L. n.v.)] & Prolus. Fl. Jap.: 239 (1867). 1.5–6(–12) cm long, 5–12-flowered. Primary – A. trisperma (Miq.) Baker in Hook., Fl. Brit. bracts broadly ovate to orbicular, 2–5 mm long, India 2(4): 181, in obs. (1876); Kitag., Neo- 1.5–4 mm wide, striate, abaxial surface glabrous Lineam. Fl. Mansh.: 393 (1979); P. Y. Fu & Y. or sparsely pubescent, ciliate, apex acute, A. Chen, Fl. Pl. Herb. Chin. Bor.-Or. 5: 163, t. obtuse, rounded, truncate or bilobed; secondary 72 (1976); W. T. Lee, Lineam. Fl. Kor. 1: 554 bracts absent. Pedicels 1–5 mm long, reflexed (1996). – A. edgeworthii var. trisperma (Miq.) pubescent. Bracteoles absent. Ohwi [Fl. Jap.: 694 (1953), n.n.] Bull. Natn. Sci. Chasmogamous flowers: Calyx tubular, (5.5–) Mus. Tokyo no. 33: 77 (1953), & Fl. Jap. ed. 7–10 mm long, 4-lobed, sparsely pubescent; Engl.: 571 (1965). tube 3.5–5 mm long, ca. 3 mm wide, adaxial A. edgeworthii var. japonica f. aidzuensis lobes broadly triangular, 2-nerved, 2–3 mm Matsuda in Bot. Mag. (Tokyo) 25: (371) (1911) long, apex entire and acuminate; lateral lobes [Syntype: Japan. Bandaisan. G. Nakahara broadly triangular, acuminate, ca. 3 mm long, 242 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue abaxial lobe triangular, acuminate 3.5–4 mm 7–10 mm across. long. Corolla white and purple-tipped or entirely Chromosome numbers: 2n = 22 (Fukui and white. Standard broadly obovate to almost Suda 1975, etc. fide Nishikawa 2008). A somatic orbicular with short claw, 12–15 mm long, 8–10 count of 2n = 20 (Yeh et al. 1983) seems to be mm wide, auriculate at base of lamina, claw erroneous in the same reason as pointed out by ca. 2 mm long; wings oblong, 11–13 mm long, Kumar and Hymowitz (1989) for the same count lamina ca. 5 mm long, with a long stipe, minute in Amphicarpaea bracteata. auricles at base, connate with keels at a point on Distribution: Nepal, NE. India (Assam), lamina, apex obtuse; keels similar but slightly Vietnam, China, Taiwan, Japan (Kyushu, shorter than wings, connate long abaxial margin, Shikoku, Honshu, Hokkaido, and S. Kuriles), apex obtuse. Vexillary stamen entirely free, 11– Korea, and Far East Russia (Khabarovsk and 12 mm long, others alternately slightly longer or Primorye). shorter, as long as vexillary one. Pistil slightly A close affinity in external morphology of longer than androecium, stipitate, stipe ca. 2 mm Amphicarpaea bracteata and A. edgeworthii long, ovary 2–2.5 mm long, style long filiform, was observed by Turner and Fearing (1965) pubescent on lower half, slightly thickened and and both were treated as geographic subspecies glabrous on upper half, 6–8 mm long. (Ohashi 1966). Pollen structure differs between Inflorescences with aerial cleistogamous the two species: triporate or tetraporate (Meeson flowers axillary at the same node with 1974). They were indicated as distinct in the chasmogamous flowered-inflorescence, raceme, phylogenetic analyses (Lee and Hymowitz 2001, 0–2 cm long, 2–4-flowered, or axillary at leaf- Parker et al. 2004). node, 1-flowered. Aerial cleistogamous flowers enclosed within calyx, calyx similar to that of 3. Amphicarpaea ferruginea Benth. in Miq., chasmogamous flower but smaller, 3.5–5.5 mm Pl. Jungh.: 231 (1852) [Lectotype (designated long, without petals, without stamens, pistil here): Nepalia. Wallich, Cat. n. 5514 stipitate, ovary elliptic, 1.5–2 mm long, glabrous, (K001121141) ut Glycine ferruginea Wall. (Fig. densely ciliate, with rudimental style and stigma 8)]; H. Ohashi in Fl. E. Himal. 3: 55 (1975), at apex, style reflexed along adaxial suture. in H. Hara & Williams, Enum. Fl. Pl. Nepal 2: Pods from aerial flowers narrowly oblong, 105 (1979); Sa & Gilbert in Fl. China 10: 249 shortly apiculate with style-base, compressed, (2010). – Pueraria ferruginea (Benth.) Kurz in not septate, (2–)2.5–3 cm long, 6–8 mm J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 42(4): 232 wide, glabrous and faintly reticulate veined (1874). – Shuteria ferruginea (Benth.) Baker in on lateral surfaces, densely ciliate on sutures, Hook. f., Fl. India 2: 182 (1876). 2–3(–4)-seeded; pods from aerial cleistogamous [Glycine cylindriflora Wall., Numer. List. n. flowers similar to those from chasmogamous 5516 (1831–32), nom. nud.]. flowers. Seeds from aerial pods compressed, Amphicarpaea edgeworthii Benth. var. round, 3.5–4 mm across, dark brown with black rufescens Franch., Pl. Delavay.: 178 (1890) spots, hilum short, ca. 1 mm long, rim-arillate. [Syntype: China. Yunnan. Delavay 3232, 3267 Subterranean flowers cleistogamous, & 3693 (all P)]. – A. rufescens (Franch.) [Thuan, globular, enclosed within calyx, white, ca. 1 Adansonia, n.s. ser. 2, 12(2): 304 (1972), comb. mm long, apetalous, rudimental stamens 10, nud.] Y. T. Wei & S. K. Lee in Guihaia 5(3): 174 ovary 1, style recurved, minute, stigma minute, (1985). rudimental. Subterranean pods subglobose, Shuteria ferruginea var. bracteosa Prain 1–2-seeded, pericarp pale pink, glabrous, with a in J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: reflexed style. Seeds pale pink with black spots, 403 (1897) [Syntype: Sikkim. Clarke 13493; December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 243

Fig. 8. Lectotype of Amphicarpaea ferruginea Benth. K001121141 (©The Board of Trustees of the RBG, Kew). 244 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Fig. 9. One of the holotype of Shuteria africana Hook. f. K000087082 (©The Board of Trustees of the RBG, Kew). December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 245

Fig. 10. Syntype of Shuteria hirsuta Baker, K000894837 (©The Board of Trustees of the RBG, Kew). 246 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Khasia. G. Mann, Collett, Clarke 40383 (all in stamens; pedicels 0–2 mm long. Chasmogamous K)]. – A. ferruginea var. bracteosa (Prain) H. flowers 15–20 mm long; calyx tubular, 7–9 mm Ohashi & Tateishi in J. Jap. Bot. 57(1): 4, fig. 1 long, 4-lobed, sericeous; tube 5–6 mm long, (1982). adaxial lobe broadly triangular, 2–3 mm long, Shuteria anomala Pamp. in Nuovo Giorn. lateral lobes triangular, 2–3 mm long, apex Bot. Ital. ser. 2, 17: 29 (1910) [Type: China. acuminate, abaxial one slightly longer than Yunnan. Ravins des montagnes. Oct. 1904. P. laterals. Corolla blue to purple or white and Maire 178]. purple-tipped; standard obovate-elliptic, 15–18 A. ferruginea f. pauciflora Pamp. in Nuovo mm long, 8–9 mm wide, shortly auriculate, Giorn. Bot. Ital. ser. 2, 17: 30 (1910). narrowed to stipe; wings elliptic, base with long Shuteria siamensis Craib in Bull. Misc. stipes (9–12 mm long), lamina 5–6 mm long, Inform. Kew 1927: 61 (1927) [Holotype: 3–3.5 mm wide, with acute auricles at base, apex Thailand. A. F. G. Kerr 6305. (K000883274)]. obtuse; connate with keels at center of lamina; Perennial herbs with long roots. Stems keels slightly shorter than wings, 15–18 mm densely or uniformly reflexed yellowish long (stipe 10–11 mm long, lamina obliquely pubescent or ferruginous villous. Stipules broad-ovate, 4–5 × 4–5 mm), connate along deciduous, narrowly ovate to oblong, 5–8 mm abaxial to upper adaxial margin, apex obtuse. long, striate, abaxial surface appressed brown Pistil linear, stipe 3–4 mm long, ovary 4-ovuled; sericeous, apex acute. Petioles 4–12 cm long, style ca. 9 mm long, glabrous; stigma capitate. reflexed brown pubescent; stipels small, linear, Pods from chasmogamous flowers falcate, 2–3 ca. 2 mm long. Leaflets papery to thickly × 0.6–0.9 cm, stipitate, lateral surfaces densely papery, ovate or ovate-elliptic to broadly ovate, clothed with yellowish-brown pubescence, both apex attenuate to cuspidate, abaxial surfaces sutures thickened, densely ascending hairy, base densely yellowish brown villous especially on acuminate to stipe, apex beaked, straight with nerves, lateral veins prominent, 5–6 on each base of style; pods from cleistogamous flowers side, adaxial surface rather densely villous, base similar to those from chasmogamous flowers. rounded or broadly cuneate; terminal leaflet Seeds 2–3, blackish gray, reniform, ca. 5 × 3 3.5–8.5 × 2–6 cm; lateral leaflets similar to the mm. terminal one, conspicuously oblique at base. Distribution: Nepal, Sikkim, India (Assam: Pseudoracemes 2–7 cm long, simple, rarely Khasi Hills), Bhutan, Myanmar, Thailand, and branched, with 6–12-chasmogamous flowers, China (Yunnan and Sichuan). axis densely patent to reflexed brown villose. This species is similar to Amphicarpaea Primary bracts early deciduous or persistent, africana in its perennial habit and lack of ovate to broadly ovate to orbicular or depressed subterranean pods, but differs from it clearly in ovate, 3–9 mm long, 2–10 mm wide, densely the structure of apertures of pollen grains. On ciliate, apex rounded or acute to acuminate; the other hand, the structure of A. ferruginea secondary bracts absent, very rarely present, is common with those of A. bracteata and A. filiform, 2–3 mm long. Pedicels 2–6 mm edgeworthii, although these are annual with long, reflexed pubescent. Bracteoles absent, subterranean pods. very rarely persistent, narrowly ovate, ca. 2 In the protologue, Wallich, Cat. n. 5514 mm long, acute, sparsely puberulent, cilate. Glycine ferruginea Wall. and n. 5516 Glycine Cleistogamous flowers sometimes present, cylindriflora Wall. were cited. The two specimens similar to undeveloped chasmogamous flowers, are referable to Amphicarpaea ferruginea. The usually 2 at the base of pseudoracemes, enclosed former one is selected for the lectotype of the in calyx 2–5 mm long, with shorter pistil than name. December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 247

Species excluded from Amphicarpaea (ICN 52.1: McNeill & al. 2012, Ohashi & al. 1. Shuteria africana Hook. f. in J. Proc. 2014). Linn. Soc., Bot. 7: 190 (1864) [Holotype (2 Shuteria anabaptis C. Y. Wu in J. W. China sheets): “Cameroon Mountains, alt. 7000 ft. (Fl. Border Res. Soc. 16: 173 (1946), in adnot. Nov.)”, Y. Mann Nov. 1974 (K000087082 [sheet Amphicarpaea lineata Chun & T. C. Chen 1/2]) (Fig. 9) and (K000087083 [sheet 2/2])]. in Acta Phytotax. Sin. 7: 23, pl. 8 (1958) [Type: – Amphicarpaea africana (Hook. f.) Harms in CHINA. Hainan. Paoting Hsien. 6 Jan. 1934. H. Repert. Spec. Nov. Regni Veg. 17: 136 (1921), Y. Liang 64056. PE–holotype; ISBC–isotype]; in adnot.; Hauman in Fl. Congo Belge 6: 90, t. Sa and Gilbert in Fl. China 10: 249 (2010). 7 (1954); Turner & Fearing in Southwest. Nat. Thuan (1972) noted that the types of Shuteria 9: 211 (1965); Verdc. in Fl. Trop. E. Africa, hirsuta Bak. (Type: Mt. Khasia, Hooker f. & Legum. (Part 4) 511, fig. 73 (1971); H. Ohashi, Thomson s.n. K) and S. ferruginea Bak. (Type: Nemoto & Onodera in J. Jpn. Bot. 80: 133, fig. Nepal, Graham in herb. Wallich 5514. K) are 157 pollen (2005). conspecific. Although Shuteria ferruginea was This species is returned to Shuteria based not cited as a synonym of S. hirsuta in his paper, mainly on its structural similarity in pollen he cited the type of S. ferruginea ‘Graham in to other members in the genus. The pollen of herb. Wallich 5514 (K)’ under S. hirsuta. Amphicarpaea has no colpori but linear grooves or slight colporus-like furrows. The membrane We are grateful to Dr. Y. Tateishi and Ms. J. on the furrows has sculpture of quite similar to Iketsu in reference to their unpublished works that of the mesocolpium. On the other hand, performed in 1986–1988 at the Plant Taxonomy Shuteria has clearly developed colpori with Laboratory in The Biological Institute of margines and the colporus membrane is smooth Tohoku University, to Prof. T. Nemoto of or verrucate. Ishinomaki Senshu University for comments, There are two sheets of the holotype of to Dr. K. N. Gandhi of Harvard University Shuteria africana Hook. f. in Kew: K000087082 Herbaria for nomenclature of the generic name [sheet 1/2] and K000087083 [sheet 2/2]. The Amphicarpaea, and to Ms. Emily Wood of species was described with the flower pedicels Harvard University Herbaria for reading a and the calyx in the protologue, but these organs preliminary manuscript. The figures of the type are found on K000087082. specimens are reproduced with the consent of the Royal Botanic Gardens, Kew (©The Board 2. Shuteria hirsuta Baker in Hook. f., Fl. of Trustees of the Royal Botanic Gardens, Kew) Brit. India 2: 182 (1876) [Type: INDIA. Khasia. for this study. We thank Dr. G. Lewis and Ms. Regio temp., alt. 3–5000 ped. J. D. Hooker & T. Alicia Musson of Royal Botanic Garden for Thomson s.n. (K–lectotype designated by Thuan reproducing the images. in 1972); Sikkim. Regio subtrop., alt. 3–5000 ped. J. D. Hooker (K000894837–syntype) (Fig. References 10)]; Thuan in Adansonia ser. 2, 12: 303 (1972), Bentham G. 1865. Leguminosae. In: Bentham G. and Hooker J. D., Genera Plantarum 1: 434–600. Reeve & excl. ‘Glycine ferruginea Grah. (nom. nud.)’. Co., London. Pueraria anabaptista Kurz in J. Asiat. Soc. Erdtman G. 1960. The acetolysis method. A revised Bengal, Pt. 2, Nat. Hist. 45(4): 253 (1877, not description. Svensk. Bot. Tidskr 54: 561–564. 1876) [Type: not designated. Habitat Pegu Espert S. M., Sede S. M., Ruiz L. K. Fortunato R. H. and Yomah; Ava; Khakyen Hills.]. Shuteria hirsuta Poggio L. 2008. New chromosome reports in the subtribes Diocleinae and Glycininae (Phaseoleae: Baker was cited as the synonym under the name, Papilionoideae: Fabaceae). Bot. J. Linn. Soc. 158: hence Pueraria anabaptista is superfluous name 336–341. 248 The Journal of Japanese Botany Vol. 91 Centennial Memorial Issue

Ferguson I. K. and Skvarla J. J. 1981. The pollen Nicolson D., Greuter W., Demoulin V. and Brummitt R. morphology of the subfamily Papilionoideae 1984. On changes made in Appendix III (Sydney (Leguminosae). In: Polhill R. M. and Raven P. H. Code). Taxon 33: 310–316. (eds.), Advances in Legume Systematics. Part 2, pp. Nishikawa T. (ed.) 2008. Chromosome atlas of flowering 427–463. Royal Botanic Gardens, Kew. plants in Japan. National Museum of Nature and Fukui J. and Suda Y. 1975. Chromosome numbers of Science Monographs No. 37. National Museum of Japanese Amphicarpaea Ell. J. Fac. Agric. Iwate Univ. Nature and Science, Tokyo. 12: 317–320. Nuttall T. 1818. The genera of North American plants, and Guo X. M., Qiao Y. K., Wang Y. X., Liu Y. J. and Liu J. a catalogue of the species to the year 1817. vol. 2. D. Z. 2003. Observation on pollen and leaf epidermis of Heartt, Philadelphia. Amphicarpaea edgeworthii by SEM. Acta Bot. Boreal.- Ohashi H. 1966. Leguminosae. In: Hara H. (compil.), The Occident. Sin. 23: 2072–2076. Flora of Eastern Himalaya. pp. 135–166. University of Hesse M., Halbritter H., Weber M., Buchner R., Frosch- Tokyo, Tokyo. Radivo A. and Ulrich S. 2009. Pollen terminology, An Ohashi H. 1975. Leguminosae. In: Ohashi H. (compil.), illustrated handbook. Springer-Verlag, Wien. The Flora of Eastern Himalaya 3: 54–72. University of Hoen P. 1999. Glossary of pollen and spore terminology. Tokyo Press, Tokyo. 2nd and rev. ed. Laboratory of Palaeobotany and Ohashi H. and Mill R. R. 2000. Hylodesmum, a new name Palynology, Utrecht University, Utrecht. for Podocarpium (Leguminosae). Edinb. J. Bot. 57: Huang T. C. 1968. Pollen grains of Formosan plants (4). 171–188. Leguminosae. Taiwania 14(2): 154–196. Ohashi H., Nagamasu H. and Murata J. (eds.) 2014. Huang T. C. 1972. Pollen Flora of Taiwan. National Taiwan International Code of Nomenclature for algae, fungi, University Botany Department Press, Taipei. and plants (Melbourne Code). Japanese Edition. Hutchinson J. 1964. Genera of Flowering Plants Hokuryukan, Tokyo. (Angiospermae) based principally on the Genera Ohashi H., Nemoto T. and Onodera R. 2005. Pollen Plantarum of G. Bentham and J. D. Hooker. vol. 1. morphology of the Japanese Phaseoleae (Leguminosae: Oxford University Press, London. Papilionoideae). J. Jpn. Bot. 80: 125–160. Ikuse M. 1956. Pollen Grains of Japan. Hirokawa Parker M. A., Doyle J. L. and Doyle J. J. 2004. Comparative Publishing Co., Tokyo (in Japanese). phylogeography of Amphicarpaea legumes and their Kumar P. S. and Hymowitz T. 1989. Where are the diploid root-nodule symbionts in Japan and North America. J. (2n = 2x = 20) genome donors of Glycine Willd. Biogeography 31: 425–434. (Leguminosae, Papilionoideae)? Euphytica 40: 221– Punt W., Blackmore S., Nilsson S. and Thomas A. Le, 226. 1994. Glossary of pollen and spore terminology. Lackey J. A. 1981. Phaseoleae DC. (1825). In: Polhill Laboratory of Palaeobotany and Palynology, LPP R. M. and Raven P. H. (eds.), Advances in Legume Contributions Series, No. 1. LPP Foundation, Utrecht. Systematics. Part 1. pp. 301–327. Royal Botanic Sa R. and Gilbert M. G. 2010. Amphicarpaea. In: Wu Z. Gardens, Kew. G., Raven P. H. and Hong D. Y. (eds.), Flora of China Lee J. and Hymowitz T. 2001. A molecular phylogenetic 10: 249–250. Science Press, Beijing and Missouri study of the subtribe Glycininae (Leguminosae) derived Botanical Garden Press, St. Louis. from the chloroplast DNA rps16 intron sequences. Sa R., Wu T. Chen D., Zhang D., Sun H., Huang P., Gilbert Amer. J. Bot. 88: 2064–2073. M. G. Thulin M., Wilmot-Dear M. and Ohashi H. Liu B. 1991. Pollen morphology of Phaseoleae in China 2010. 19. Tribe Phaseoleae. In: Wu Z. G., Raven P. H. and its significance to the art. Bot. Research (Inst. Bot. and Hong D. Y. (eds.), Flora of China 10: 196–261. Acad. sinica) (5): 59–86. Science Press, Beijing and Missouri Botanical Garden Löve Á. and Löve D. 1982. Reports. In: Löve Á (ed.), Press, St. Louis. IOPB chromosome number Reports LXXV. Taxon 31: Schively A. F. 1897. Contributions to the life history of 344–360. Amphicarpaea Monoica. Penn. Univ. Contr. Bot. Lab. McNeill J. (Chairman), Barrie F. R., Buck W. R., Demoulin 1: 270–363. V., Greuter W., Hawksworth D. L., Herendeen P. Schrire B. D. 2005. Tribe Phaseoleae. In: Lewis G., Schrire S., Knapp S., Marhold D., Prado J., Prud’homme B., Mackinder B. and Lock M. (eds.), Legumes of the van Reine W. F., Smith G. F., Wiersema J. H. and World. pp. 393–431. Royal Botanic Gardens, Kew. Turland N. J. (Secretary) 2012. International Code of Stefanović S., Pfeil B. E., Palmer J. D. and Doyle J. J. Nomenclature for algae, fungi, and plants (Melbourne 2009. Relationships among Phaseoloid Legumes based Code). Koeltz Scientific Books, Koenigstein. on sequences from eight chloroplast regions. Syst. Bot. Meeson B. W. 1974. Pollen morphology of Amphicarpaea 34(1): 115–128 (Leguminosae: Phaseoleae). Sida 5: 338–340. Taubert P. 1894. Leguminosae. In: Engler A. and Prantl K., December 2016 Ohashi & Ohashi: Revision of Amphicarpaea 249

Die natürlichen Pflanzenfamilien. Teil III, Abteilung3 : of the genus Amphicarpaea (Leguminosae). The 70–96. Engelman, Leipzig. Southwest. Naturalist 9: 207–218. Thuan N. van 1972. Révision du genre Shuteria Yeh M., Maekawa F. and Yuasa H. 1983. Chromosome (Papilionaceae). Adansonia, sér. 2, 12: 291–305. numbers of tribe Phaseoleae, Leguminosae. Res. Inst. Turner B. L. and Fearing O. S. 1965. A taxonomic study Evol. Biol. Sci. Rep. 2: 37–44.

a b 大橋広好，大橋一晶：マメ科ヤブマメ属の分類，特に花粉形態に基づく Shuteria との関連ヤブマメ属はマメ亜科インゲンマメ連ダイズ亜 Chun & T. C. Chen は，2010 年の Flora of China 10 で独連 subtribe Glycininae に含まれる (Schrire 2005)．立種と認められているが，タイプ標本をも精査してみて， Turner and Fearing (1965) はヤブマメ属をアフリカの Shuteria hirsuta Bakerに他ならないことが明らかとなった． Amphicarpaea africana，アジアの A. edgeworthii，アメその結果，ヤブマメ属をアジアの A. edgeworthii Benth.とリカの A. bracteata の 3 種に整理した．その後，アジア A. ferruginea Benth.，およびアメリカの A. bracteata (L.) から A. ferruginea, A. lineata などが加えられ，最近では Fernald の 3 種に整理した．Ohashi (1966) はヤブマメを形 4–5 種よりなるとされている (Schrire 2005)．ヤブマメ属態的な類似からアメリカ産 A. bracteata の地理的亜種とし Amphicarpaea は形態的には Shuteria に最も近縁とされてていたが，花粉形態の違い (Meeson 1974) と分子系統解いるが，分子系統解析の結果からかけ離れたものである析の結果 (Lee and Hymowitz 2001, Stefanović et al. 2009) ことが分かった (Lee and Hymowitz 2001, Stefanović et al に基づいて本論文で別種に戻した．ヤブマメの学名は 2009)．ヤブマメ属と Shuteria は外部形態では区別が困難 Amphicarpaea edgeworthii Benth. であり，その異名とそれである一方，花粉形態では発芽溝の発達程度に明瞭な違らのタイプ標本を整理した．また，ヤブマメの開放花，閉いがある．本研究ではヤブマメ属と Shuteria の花粉形態鎖花，地上生豆果，地下生豆果について詳しく記載した．の比較によって，A. africana (Hook. f.) Harms は Shuteria （a東北大学植物園津田記念館，に属することを明らかにした．また，Amphicarpaea lineata b岩手医科大学薬学部）