hosta tsushimensis - hosta library · suggest h. tsushimensis migrated from korea to tsushima...

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Hosta Species UpdateThe Hosta LibraryORG20060926©W. George Schmid 2010 Original Edition 20060926 Revised Edition 20101220 R1 H. tsushimensis Fujita 1976 H. tsushimensis N.Fujita f. albiflora (S.Toyama) N.Fujita Acta Phytotaxonomica et Geobotanica, Vol. 27, 3/4:91, ic. 9 1976. ツシマギボウシ = 対馬擬宝珠 = Tsushima Gibōshi シロバナツシマギボウシ = H. tsushimensis f. albiflora = Shirobana Tsushima Gibōshi History & Nomenclature: In Japan this species is called Tsushima Gibōshi, the “hosta from Tsushima,” (Tsushima Island). In 1976, the ecologist N. Fujita de- scribed H. tsushimensis from popu- lations found in various locations on both of the large Tsushima Islands. Its main habit is located on Tsu- shima-shichō (対馬支庁) on Mount Izuhara ( 厳原 ) and on Shimono- shima ( 下島 ), on Mount Ariake- yama (有明山). M.G. Chung located colonies on Mount Izuhara and he considered only material collected on Tsushima Island in his 1990 study. This makes it difficult to cor- relate it to two other related taxa: H. jonesii and H. tibae. Fujita and Tamura (2008) considered H. tibae a variety of H. tsushimensis. rt of Habitat and Biology: The habitat of this species is located in the Tsushima Strait (対馬海峡 = Tsushima Kaikyō), which is the eastern channel of the Korea Strait. The Tsushima Strait lies between Korea and Japan, connecting the Sea of Japan (East Sea) and the East China Sea. The two main islands are called Kami (north) and Shimo (south) and there are many small islands, all considered pa 2010-12-20 - 1 - H. tsushimensis N. Fujita (cult. voucher) Nagasaki-ken (長崎県);Tsushima-shichō (対馬支庁); Izuhara (厳原); Vouch.: Chung 1614

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Page 1: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

Hosta Species Update●The Hosta Library●ORG20060926●©W. George Schmid 2010 Original Edition 20060926 Revised Edition 20101220 R1

H. tsushimensis Fujita 1976 H. tsushimensis N.Fujita f. albiflora (S.Toyama) N.Fujita

Acta Phytotaxonomica et Geobotanica, Vol. 27, 3/4:91, ic. 9 1976. ツシマギボウシ = 対馬擬宝珠 = Tsushima Gibōshi

シロバナツシマギボウシ = H. tsushimensis f. albiflora = Shirobana Tsushima Gibōshi

History & Nomenclature: In Japan this species is called Tsushima Gibōshi, the “hosta from Tsushima,” (Tsushima Island). In 1976, the ecologist N. Fujita de-scribed H. tsushimensis from popu-lations found in various locations on both of the large Tsushima Islands. Its main habit is located on Tsu-shima-shichō (対馬支庁) on Mount Izuhara (厳原 ) and on Shimono-shima (下島 ), on Mount Ariake-yama (有明山). M.G. Chung located colonies on Mount Izuhara and he considered only material collected on Tsushima Island in his 1990 study. This makes it difficult to cor-relate it to two other related taxa: H. jonesii and H. tibae. Fujita and Tamura (2008) considered H. tibae a variety of H. tsushimensis.

rt of

Habitat and Biology: The habitat of this species is located in the Tsushima Strait (対馬海峡 = Tsushima Kaikyō), which is the eastern channel of the Korea Strait. The Tsushima Strait lies between Korea and Japan, connecting the Sea of Japan (East Sea) and the East China Sea. The two main islands are called Kami (north) and Shimo (south) and there are many small islands, all considered pa

2010-12-20 - 1 -

H. tsushimensis N. Fujita (cult. voucher)Nagasaki-ken (長崎県);Tsushima-shichō

(対馬支庁); Izuhara (厳原); Vouch.: Chung 1614

Page 2: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

KOREA Location Map of Tsushima Island

Sea of Japan

Place of Type Collection Mt. Ariake (Shown by ♦ Arrow; No. 3037 KYO)

◄Tsushima Islands

♦ ← Korea Strait JAPAN (Kyushu) Del. W.G. Schmid 1991 Nagasaki-ken ( 長 崎 県 ). H. tsushimensis grows in various locations on the main islands. This includes dry, higher ele-vations wooded areas and un-der subshrubs. Other popula-tions are found in bottom land, which contains more moisture, near streams or near the coast. This species is very polymor-phic and many different leaf shapes can be found from narrow-lanceolate to about ovate. Some have straight mar-gins, others have wavy edges. A number of variegated mutants found in the wild have been se-lected and named. As with H. jonesii and H. tibae, H. tsu-shimensis also develop branch-ing racemes in some popula-tions. Its flowers are lighter in color and whitish purple on the outside of the perianth. It is similar to H. tardiva but differs by its later later flowering time,

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HAriake

Hosta

. tsushimensis (cult. voucher) -yama (有明山); Shimono-shima (下島)

NA Voucher 45167 Hill R.G. © W.G. Schmid 1989.04.17

-

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the

its wider, more glossy and cor-date leaves with smooth veins underneath, and its good fertility. A white-flowered form is known as H. tsushimensis f. albiflora (S. Toyama) Yonek., comb. nov. It is rare in the wild habitat. J. Jap. Bot., 80:6 (2005). H. tsushimensis, H. jonesii, and H. tibae form a closely related group but Chung (1990) did not include the latter in his studies. Ohba, et. al., (1987) suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together with other glacial remnants. My studies show, it is also possible that the reverse took

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H. tsushimensis (cult. voucher 45167) Ariake-yama (有明山); Shimono-shima (下島) Clone No. 3/Hosta Hill R.G. © W.G. Schmid

H. tsushimensis N. Fujita 1976 (in situ) Nagasaki-ken (長崎県);Tsushima-shichō; (対馬支庁); Izuhara (厳原)

Note: Species growing in their natural habitat often grow under difficult conditioncompeting with other plants. The result is seen in this photo. Compare to photos of

this species cultivated in gardens. (Follow arrow for flower buds)

Page 4: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

place and that H. jonesii and H. tsushimensis orig-inated with the ancestral populations of H. tibae in the northwestern Kyushu habitat. Chung (1990) argues that H. tsushimensis and H. jonesii may have developed from elements of H. minor, separated by the last glacial event but it seems more likely that the propagule was a species from Japan, based on RAPD analysis (banding patterns) and distribution patterns. H. tsushimensis and H. jonesii are morpho-logically much closer to the Japanese populations than the highly differentiated native Korean species: H. yingeri, H. laevigata, H. clausa, H. minor, H. venusta and H. capitata. Although scape branching has been observed in many species, it is not a consistent trait but the natural populations of H. jonesii, H. tsushimensis and H. tibae include many clones exhibiting consistent branching of the scapes which may be an evolutionary response to a lack of compatible pollinators in a windswept insular and coastal environment. Specimens (in L) collected by von Siebold in 1827 also show this branching (in H. tibae) [see Species Update; H. tibae, pages 4 and 5]. These species are related and form a group, which has its largest representation on Kyushu. Plant Morphology: Plant size 45 cm dia. by 20 cm high (18 by 8 in.). Petiole 17.5–25 cm by 6–.8 cm wide (7–18 by 0.25–0.35 in. wide), slightly winged, erect, green, with purple dots. Leaf 15–22.5 cm by 7.5–12.5 cm wide (6–9 by 3–5 in.), erect, ovate to ovate-cordate, tip acuminate, slightly undulate, wavy in the margin, smooth, light green above, glossy lighter green below. Venation 5–9, sunken above, projected, smooth below. Scape 65–90 cm long (26–30 in.), erect, obliquely ascending, lax, green, purplish red dot-ted. Sterile bracts, 2–3, clasping the stem; fertile bracts, navicular, long, thick, grooved, whitish-green, persisting, not withering at anthesis. Raceme 25–30 cm (10–12 in.), loosely arranged, to 30 flowers, frequently branching. Flowers 4–5 cm long and 4 cm broad (1.5–2 by 1.50 in.), translucent lines end at center, whitish to purple outside, inner nerves purple, tepals color: ▲▲(Type B in W.G. Schmid;

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H. tsushimensis (cult. NA voucher 45167 clone 3) Dr. S. B. Jones Garden - © W.G. Schmid 1987.07.18

Page 5: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

1991) perianth expanding, funnel-shaped, lobes spreading acutely; pedicels to 1.5 cm (0.5 in.) white, purple-dotted, stamens projecting greatly beyond perianth. Anthers whitish posterior, sparsely purple dotted. Late August/September. Fertile.

Karyotype-Chromosomes: Sporophytic Count = 60; 12 large, 48 small; (2n). Pollen: (Pollen shape after Erdtman, 1966): H. tsushimensis = Subtype RG(IIA) (rugulate granulate): OS (oblate-spheroidal); size P 69.0 ± 1.9 × E 64.3 ± 2.4 (Sizes given in µm ±2–10% polar axis (P) × equatorial axis (E)). As typified by M.G. Chung and S.B. Jones, 1989;

jonesii, the latter having

lated to H. jo

Genome Size: DNA content (2C) in pg (one (10-12) gram) = 17.3 ± 0.12. (Zonneveld, B.J.M. and F. Van Iren (2001); with other data included, this value indicates close relationship with H.a pg of 17.5 ± 0.09)

DNA Banding: Recent RAPD analysis (Y. Yu, and W.G. Schmid. 2005). In additions to otherrelated species accessions (See Fig. B) were compspecies shown in the banding pattern were compa(5'-GTTTCGCTCC-3'), generated three bands asband (850bp) common to (36) H. tardiva, (39) H. tthe separation of (41) H. tsushimensis from the which was common to (36) H. tardiva and (40) Hfrom (39) H. takahashii. The third band (676bp) atibae from (36) H. tardiva. Separation of H. tsuindication that this species is closer re 36 39 40 41

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H

. tsushimensis: Pollen TypeRG IIA Grain Surface Detail

SEM × 4000 (M.G. Chung)

nesii (confirmed by DNA content).

2002; Sauve, R.J., S. Zhou, Y. Yu, taxa, the banding patterns of 4 ared in the 2002/2005 study. The 4 red using a single primer OPB-01 shown in Fig. B. The polymorphic akahashii, (40) H. tibae allowed for group. The second band (937bp), . tibae allowed for their separation llowed for the separation of (40) H. shimensis from this group is an

Banding Pattern (Fig. B)

36 = H. tardiva 39 = H. takahashii 40 = H. tibae 41 = H. tsushimensis

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201

H. tsushimensis Holotype

Ariake-yama (有明山); Shimono-shima (下島); Tsushima-shichō (対馬支庁)Nagasaki-ken (長崎県); Kyūshū (九州)

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Holotp. KYO No. 3037. Kyoto Univ. Herbarium ♦ 京都大学総合博物館

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H. tsushimensisIsotypus: TNS 249544 (F. 301);

Coll. loc. cit.: Ariake-yama (有明山); Sh(対馬支庁); Nagasaki-ken

Isotype (01218) H. Koyama #3037; 1969.08.22. imono-shima (下島); Tsushima-shichō

(長崎県); Kyūshū (九州).

-

Page 8: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

Taxonomic Type and Synonymy: H. tsushimensis Fujita. Acta Phytotaxonomica et Geobotanica, Vol. 27, 3/4:91, ic. 9 (1976). W.G.

Schmid, The genus Hosta: Gibōshi Zoku (ギボウシ属): 113, 325, f. 3-76, t. 77 (1991). [Note: H. tsushimensis var. tsushimensis N. Fujita et M. N. Tamura: Acta Phytotaxonomica et Geobotanica, Vol. 59(1): 31-36(2008). The stat. nov. of H. tibae as a variety of H. tsushimensis, namely H. tsushimensis var. tibae proposed by the authors is not accepted here (see H. tibae)].

Type: Holotype in KYO, No. 3037. Loc cit. coll.: H. Koyama, Ariake-yama (有明山); Shimono-shima (下島), Tsushima-shichō (対馬支庁), Nagasaki-ken (長崎県); Kyūshū (九州); (see page 7). Isotype in TNS 249544, H. Koyama (page 8).

Hab.: Insular climate habitat includes dry, higher elevations wooded areas and under sub-shrubs; also bottom land containing more moisture on Tsushima-shichō (対馬支庁), Nagasaki-ken (長崎県); Kyūshū (九州);

H. tsushimensis f. albiflora (S. Toyama) K. Yonek., Basionym: Hosta minor f. albiflora S.Toyama in J. Jap. Botany 36:141 (1961).

comb. nov. J. Jap. Bot. 80:6 (2005). Type: S. Toyama, 1932.09.06 Herb. Biol. Inst., Fac. Lib. Arts, Nagasaki Univ.

Nagasaki; loc cit. coll.: Ariake-yama ( 有明山 ); Shimono-shima ( 下島 ), Tsushima-shichō (対馬支庁), Nagasaki-ken (長崎県); Kyūshū (九州);

Hab.: Insular climate habitat includes dry, higher elevations wooded areas and under sub-shrubs; also bottom land containing more moisture on Tsushima-shichō (対馬支庁), Nagasaki-ken (長崎県); Kyūshū (九州);

Japanese Language Synonyms: ツシマギボウシ = Tsushima Gibōshi = H. tsushimensis シ ロ バ ナ ツ シ マ ギ ボ ウ シ = Shirobana

Tsushima Gibōshi = H. tsushimensis f. albiflora.

ral Names: t

sushima Hosta

HorticultuH. minor incorrecT H. tsushimensis (cult. voucher) ►►► Flower Detail (NA 45167) Hosta Hill R.G. © W.G. Schmid

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H. tsushimensis in Cultivation: This species is infrequently seen in gardens. There are a number of sports and var-iegated hybrids grown in gardens, but the species is not popular. There are a few plants sold that are incor-rectly identified but most are the “true” species. As seen in the in situ photo-graph on page 3, in culti-vation this species is more vigorous in growth and is an attractive horticultural sub-ject. Of high garden interest are the branched scapes seen in some of the cultivated clones. Branched scapes are also observed in H. tibae, which grows in Nagasaki Prefecture directly across the Korea Straiand in H. jonesii endemic to islands on tscapes lend an unusual look to scape eloWhether branched or not branched, the gits foliage, but rather in its flowers. In Acrown and each scape is loaded with dopurple striped inside, funnel-shaped flowe

2010-12-20 -

HAriake

Hosta

. tsushimensis (cult. voucher) -yama (有明山); Shimono-shima (下島)

NA Voucher 45167 Hill R.G. © W.G. Schmid 1989.04.17

t and Tsushima Island located in the strait he southern cast of Korea. The branched ngation and while the species is in flower. arden worthiness of this species lies not in ugust, numerous scapes rise from the leaf zens of purple, white striped outside, and rs.

◄ H.

tsushimensis (cult.)

©T. Avent; Plant Delights

Nursery

9 -

Page 10: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

H. tsushimensis (cult. voucher) (NA Voucher 45167) ▲ Detail of bud initial ► Elongating scape/raceme w/ bud initials Hosta Hill R.G. © W.G. Schmid 1989.08.22 H. tsushimensis Detail of bud initial ► Court.: © yahoo.co.jp-giboushiblog

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Page 11: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

H. tsushimensis f. albiflora (in situ) ►►► Ariake-yama (有明山) Shimono-shima (下島) In dry area along road flowers white Court. : © 対馬植物図鑑

H. tsushimensis (cult. voucher)

NA Voucher 45167 Under cultivated conditions, the scapes become longer than those found in situ. Compare in situ photograph on page 4

Hosta Hill R.G. © W.G. Schmid

1989.09.07

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Page 12: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

H. tsushimensis

(in situ Ariake-yama (有明山) Shimono-shima (下島),

Tsushima-shichō (対馬支庁) In damp area near stream.

Flower color is darker

This may be:

三色ツシマギボウシ = H.‘Sanshoku Tsushima’

Court. © 対馬植物図鑑

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H. tsushimensis (cult.) Research specimen (Court.: © yahoo.co.jp-giboushiblog)

The lighter color is attributed to shade location)

Page 13: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

Named Japanese Sports or Variants: H. ‘Ogon Tsushima’ is the correct cultivar name is a yellow form found in cultivation and possibly in the wild. The Japanese named it 黄金ツシマギボウシ = Ogon Tsushima Giboshi. It retains its bright yellow color. By some called H. tsushimensis ‘Golden’ or H. tsushimensis ‘Ogon’. Both of these names are incorrect because their formulation goes against the rules of the International Code of Nomenclature for Cultivated Plants (ICNCP).

H.‘Ogon Tsushima’ (黄金ツシマギボウシ)

Ogon Tsushima Giboshi)

Photo © B. Skaggs Hosta Library

H. ‘Sanshoku Tsushima’ is reportedly a mutation found in the wild with tri-colored flowers, dark crimson at the base of the narrow tube, and purple on the petals and has large white transparent lines. It does not come true from seed so is considered a cultivar (H. takahashii has a similar morph, which see). The Japanese named it 三色ツシマギボウシ = Sanshoku Tsushima Giboshi. H. tsushimensis ‘Sanshoku Tsushima’ hort. is an incorrect cultivar name and against the rules of the Inter-national Code of Nomenclature for Cultivated Plants (ICNCP). H. tsushimensis ‘Albiflora’ (the botanical name is H. tsushimensis f. albiflora) is the white-flowered form of H. tsushimensis and its valid botanical name was published in 2005 as H. tsushimensis f. albiflora (S. Toyama) K. Yonekura, comb. nov. (J. Jap. Bot. 80:6 (2005). In Japan and before this, it was called 白花ツシマギボウシ = Shirobana Tsushima Giboshi (= white-flowered Tsushima hosta). This form is rarely found in the wild and reportedly does not come true from seed, although I was not able to confirm that source. Notwithstanding, if H. tsushimensis f. albiflora is considered a cultivar under the ICNCP, its name must to be H. tsushimensis ‘Albiflora’ per ICNCP Chapter V, §2. 19.6, which requires that the former correct name of the taxon be retained as the cultivar epithet (in the event the taxonomic unit is reclassified as a cultivar). The following are incorrect names used: H. ‘Shirobana Tsushima’, H. tsushimensis ‘Shirobana Tsushima’, and H. tsushimensis ‘Shirobana’.

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H. tsushimensis (cult. Voucher) NA Voucher 45167 Hosta Hill R.G. © W.G. Schmid 1989.09. 21 Three-locular seed capsules ► Here on unbranched scape/racemes Three-locular seed capsules ► Here on branched scape/racemes

◄◄ H. ‘P. M. Stripe’

H. tsushimensis ‘P.M. Stripe’

Photo ©C. Brashear Hosta Library

Darker green veining on lighter green leaf

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Page 15: Hosta tsushimensis - HOSTA LIBRARY · suggest H. tsushimensis migrated from Korea to Tsushima Island and Kyushu, Japan, during the Pleistocene and speciated on the island together

Horticultural Progeny: Note: The true H. tsushimensis is not frequently seen in gardens. 2 cultivars have been listed, which involve a yellow sport of this species as a pod (♀ H. tsushimensis) parent or pollen (see below). There are several sports in cultivation. H. ‘Beet Salad’ = ♀ H. clausa normalis F2 seedling × ♂ (H. 'Ogon Tsushima' x H.

yingeri F2 seedling)] by R. Solberg (2009) H. ‘Sun Catcher’ = ♀ H. ‘Ogon Tsushima’ × ♂ H. yingeri by R. Solberg (2004) H. ‘Whiskey Sour’ = ♀ H. ‘Ogon Tsushima’ × ♂ H. yingeri by R. Solberg (2002)

◄◄ H. ‘Whiskey Sour’ R. Solberg (2002)

Photo © I. Scroggy

H. tsushimensis (cult.) Typical flower color ▲ (Court.: © yahoo.co.jp-giboushiblog) Flowers lighter on branched racemes ► Hosta Hill R.G. © W.G. Schmid 1989.09. 21

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References: Chung, M.G. and Jones, S.B. 1989. Pollen morphology of Hosta Tratt. (Funkiaceae)

and related genera. Bulletin of the Torrey Botanical Club, Vol. 116, 1:31–44. Chung, M.G. 1990. A Biosystematic Study on the genus Hosta (Liliaceae/Funkiaceae)

in Korea and Tsushima Island of Japan. U, of Georgia, Athens; including: Morphometric and isoenzyme analysis of the genus Hosta Tratt. (with S.B. Jones and JL. Hamrick) and Isoenzyme variation within and among populations of Hosta in Korea.

Chung, M.G., J.L. Hamrick, S.B. Jones, Jr., G.S. Derda. 1991. Isozyme Variation Within and Among Populations of Hosta (Liliaceae) in Korea. Systematic Botany, Vol. 16, No. 4 (Oct. - Dec., 1991), pp. 667-684

Chung, M.G. 1996. Spatial genetic structure among Korean populations of Hosta minor and H. capitata. Bot. Bull. Acad. Sin. 37: 25-30.

Davidson, R., 1990. Letter dated January 1990 sent to Dr. R. C. Olson (Private Comm.)

Erdtman, G. 1966. Pollen morphology and plant taxonomy. Angiosperms. Hafner: N. Y.

Fujita, N. 1976. The genus Hosta (Liliaceae) in Japan. Acta Phytotaxonomica et Geobotanica, Vol. 27, (3–4) pp. 66–96

International Code of Nomenclature for Cultivated Plants = ICNCP, by the Horticultural Taxonomy Group, ISHS Commission for Nomenclature and Registration. 2004 edition.

Maekawa, F. 1937. Divisiones et plantae novae generis Hostae (1). J. Japanese Botany, 13, No. 12:893–905.

Maekawa, F. 1940. The genus Hosta. J. of the Faculty of Science, Imperial University Tokyo, Section 3 Botany, Vol. 5:317–425.

Nakai, T. 1911. Flora Koreana, pars secunda (H. minor). J. of the College of Science, Imperial University Tokyo, 31:250–251.

Ohba, H., Ishii, N. and Saito, S., 1987. Biogeography of Tsushima Island. In: Report on the Natural Resource Investigation of Tsushima Island; Nature of Tsushima Island. pp. 259–271.

Ohba, H. and Midorikawa, K., 1987. Some Remarks on the Flora of Tsushima Island, NW Kyushu, Japan. In: Report on the Natural Resource Investigation of Tsushima Island; Nature of Tsushima Island. pp. 63–78.

Sauve, R.J., S. Zhou, Y. Yu, and W.G. Schmid. 2005. Random amplified polymorphic DNA (RAPD) analysis in the genus Hosta. HortScience 40(4).

Schmid, W.G. 1988. Hosta taxonomy: a revised overview. Bull. Brit. Hosta Hemerocallis Soc. 1988 (2): 25-36.

Schmid, W.G. 1991. The genus Hosta: Giboshi Zoku (ギボウシ属). London and Portland: Timber Press.

Schmid, W.G. 1991. The Korean species. (in The genus Hosta: Giboshi Zoku. London : B.T. Batsford; Portland: Timber Press.) Pp. 297-298.

Schmid, W.G. 2004. Hosta species and DNA fingerprinting. Bull. Brit. Hosta Hemerocallis Soc. 2004: 50, 59-66.

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Schmid, W.G. 2005. Hosta DNA fingerprinting. The Hosta Journal, Vol. 36 2005 (3): 69-74.

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Yonekura, K. 2005. Taxonomic Notes on Vascular Plants in Japan (H. tsushimensis f. albiflora (S. Toyama) Yonek., comb. nov.) J. Japanese Botany, 80:334-339 (2005).

Zilis, M.R. 2000. The Hosta handbook. Rochelle: Q & by Nursery, Inc. Yu, Y. 2002. Classification of hosta species and cultivars based on RAPD analysis.

TSU Graduate School (with W.G. Schmid); published in summary in Sauve, R.J., S. Zhou, Y. Yu, and W.G. Schmid. 2005 (which see).

Zonneveld, B.J.M. and F.Van Iren. 2001. Genome size and pollen viability as a taxonomic criteria: Application to the genus Hosta. Plant Biology, 3, pp. 176-185. G. Thieme Verlag: Stuttgart.

© W.George Schmid 2010: The text and illustrations are copy-righted and are available for personal reference only. Other

contributors retain their copyright of featured photographs as noted in captions. The content may not be published in printed

form without the author’s written permission. Web quote reference:

W. George Schmid: HostaLibrary.org/species/

2010-12-20 - 17 -

H. tsushimensis (in situ) An extreme case of recurving tepals – note boat-shaped pedicels

Court.: © 2005 blog.yohbo.main.jp/yohbo 2010.09.10