H. plantaginea (Lamarck) Ascherson 1863 - HOSTA LIBRARY

Hosta Species Update●The Hosta Library●ORG20071015●©W. George Schmid 2007

Original Issue ORG2007.10.15 Revised Issue REV 1/2010.01.30

H. plantaginea (Lamarck) Ascherson 1863

Botanische Zeitung, 21:53, ic. Pl. 24 1863

H. plantaginea var. plantaginea

玉簪 = ｷﾞｮｸｼﾝ = yu zan (Chinese [recommended])

棒玉簪 = ﾎﾞｳｷﾞｮｸｼﾝ = bang yu zan (Chinese Alias/Synonym)

玉春棒 = ｷﾞｮｸｼｭﾝﾎﾞｳ = yuchun bang (Chinese Alias/Synonym)

白鶴花 = ﾊｸｶｸｶ = bai he hua (Chinese Alias/Synonym)

マルバタマノカンザシ = maruba tama no kanzashi (Japanese; Kikuchi 1934)
옥잠화 = Ok-jam-hwa (Korean name [recommended])

둥근옥잠화 = Dung-geun-ok-jam-hwa (Korean synonym)
비녀옥잠화 = Bi-nyeo-ok-jam-hwa (Korean synonym)

History and Nomenclature: H. plantaginea occupies a unique position in the

genus Hosta. It is native to China and Chinese records indicate it has been cultivated

since the Han Dynasty (漢朝 ▪ 206BC-220AD). The species was once thought to have

evolved in Hebei province (河北), the general area surrounding Beijing (北京; Běijīng

= northern capital). However, the specific physiological-ecological basics for suc-
cessful reproduction point further south (see more under Habitat and Biology).

H. plantaginea was the first hosta known
on the continent of Europe, grown from
seeds sent from Macao by French Consul
Charles de Guignes before 1784. It re-
ceived taxonomic treatment and was
established a taxon by J.B. Lamarck in
1789 as Hemerocallis plantaginea (based
on mature plants grown from this seed at
the Jardin des Plantes in Paris; the
holotype is in P). In 1790, the first live
plants were imported into England by
George Hibbert (together with H.
ventricosa, which see). Due to its tropical
look, it was first grown under glass. Some
mature specimens were part of a
collection of rare, ornamental plants
Madame Josephine Bonaparte had
assembled at the gardens of
Malmaison.◄◄◄ Left is Pierre-Joseph
Redouté’s engraving of this plant with
the name Hemerocallis Nipponia (French:
Hemerocalle du Japon (Nippon) =
Japanese Day-lily). Its attractive leaves

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and fragrant flowers made it a

popular outdoor specimen

plant, so much so, it was re-

ferred to as the “Parisian

Funkia,” and was planted all

over Paris in public and pri-

vate gardens. Soon, it spread to

other areas of France, arrived

in Germany before 1800, and

was cultivated in the United

States before 1839 (Buist

1839). This species has the best

flowers in the genus and as the

illustration to right shows; it is

more than twice as large as

those in all other hosta species

are. It has flowers of white

tepals with heavy substance Flower Size Comparison
and a waxy surface. Being the
only night-bloomer with the H. plantaginea Flower in Center
flowers opening late in the H. tibae Flower (right);H. montana Flower (left)
afternoon, their fragrance is
(Left Measure is 1 inch, Right is 1 cm)

most noticeable during evening

hours. One of the first species

of the genus to reach Europe in the early 1780s, it was also one of the first to receive

formal, taxonomic treatment. Today, under the International Rules of Botanical

Nomenclature (IR, 1935), Ed. 3:144, and the International Code of Botanical

Nomenclature, (ICBN, 1952) the genus Hosta is typified by H. plantaginea. The

nomenclature and identification of the typical Chinese species is well established.

Unfortunately, the importation and establishment of first the typical species (from

1784-1790) and different

phenotypical Japanese

forms or hybrids (from

1841-1856) into Europe has

created numerous names.

Some of these names are

applied to both the typical

species and concurrently to

Japanese phenotypes. This

◄◄◄ H. plantaginea

Typical Rounded Leaves
Court.: ©zran net

マルバタマノカンザシ

Marubatamano-Kanzashi

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has created considerable con-

fusion. For a complete list of

these synonyms and aliases, see

Taxonomic Type and

Synonymy). It is recognized

some variability does exists in

the macromorphological traits

of the Chinese H. plantaginea

but the available specimens are

for the most part very distinct

and uniform. On the other

hand, live plants and her-

barium specimens of the non-

typical Japanese forms show

distinct differences. For this

reason, the typical species is

discussed in this section and

the various named pheno-

typically different forms are

covered under H. plantaginea

var. japonica. An example

specimen of the “nontypical”

H. plantaginea is No. S2355 in

MAK. The derminavit of this

specimen is H. plantaginea var.

japonica, but it may also be H.

plantaginea var. stenantha,

judging by the very narrow

corolla (for further discussion

of the differences refers to H.

plantaginea var. japonica). As SS22FF33uu5555nnkkiinnDiiaaDMMeetgg.AAt東東rr.=aaKKHBB京京Hnn.yy((dd.MMp都都HHiiplffaaal立立ll..ankkooKKntii大大rranntaaaaagoott学学giooSSniHH;;niie牧牧eeeeFFeabbrraee野野voobbbbvallaa..dd標標arrr22r.ii((00uu.本本jiiann00jmmap44館館sspo;;cconUU))hhniee..ciddcaooaaaff TT1188oo44kk44yy))oo;;
pointed out under Habitat
and Biology, the typical
species is widely cultivated in

China, Korea, Japan, and in

gardens the world over. In

China, it has been known un-

der several names that are

treated as synonyms (or aliases). The original Chinese name is composed of two

Kanji: 玉 (meaning precious stone; jade) and 簪 (meaning “ornamental hairpin”).

The decorative part of a woman’s hairpin is made often of precious jade shaped

similar to the unopened flower bud of H. plantaginea. In a much wider interpretation,

the ancient name 玉簪 now applies to hostas and transliterated stands for “hosta.” H.

plantaginea preceded all other hostas in cultivation in China, hence the application of

the name. The aliases 棒玉簪 or 玉春棒 (yuchun bang; transliterates to “rod hosta.”

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H. ‘Chingbanyuzan’ Another Chinese alias is 白鶴花

(Chinese name for a Form of H. ‘Aphrodite’) (ban yuzan = “flower ball of or-
Known in Japan as Yae Maruba namental hairpin” = the typical
Tamanokanzashi H. plantaginea; in Japanese, the
Kanji combination translates to
八重 咲きマルバタマノカンザシ “white crane flower”). Another
name having its roots in China
Photo HL by K. Terpening has found its way into western
gardens, namely H. ‘Chingban-
yuzan’. Plants imported under
this name are none other than
the typical species. Like the
species, his cultivar develops
petaloid stamens, which might
indicate it is a transitional form
sporting to H. ‘Aphrodite’. The
name H. ‘Chingbanyuzan’ may
in fact be the Chinese name for
H. ‘Aphrodite’ named by F.
Maekawa (1940). The accepted

Chinese Kanji ( 玉 簪 ) used in

Flora of China (2000) has
carried over in Japan as the

Katakana タ マ ノ カ ン ザ シ

(transliterated = tama no kan-
zashi), which means “jewel of
the hairpin.” This Japanese
name did not come into use until
1934, when the Japanese

botanist Kikuchi published on this
subject (Kikuchi; 1934). Maekawa
(1940; p. 346) stated: “It is strange that
such a remarkable garden plant [i.e., the
typical H. plantaginea: WGS], so very
widely culti-vated in China, was
overlooked by its neighbor Japan.” There
is a Japanese variety, however, which has
been cultivated in Japan for a very long
time. It was mentioned by the Japanese

scholar Ranzan Ono ( 小 野 蘭 山 ) in

Botanical classification ( 本 草 綱 目 啓 蒙 ; H. plantaginea var. japonica

1802/1803). In the 1847 revised version of
Botanical classification (Jap.: Jūtei
Honzō-Kōmoku Keimō), this Japanese
phenotype was listed under its original

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Chinese name (玉簪 = タマノカンザシ = tama no kanzashi), which created a

nomenclature problem. Noting the Chinese species had roundish leaves, while the
Japanese had elongated leaves, Maekawa named the former maruba tama no

kanzashi (= マルバタマノカンザシ). Maruba is transliterated from the Kanji 丸葉

(= マルバ = meaning circle; rounded ─ to indicate the rounded leaves of the typical

Chinese species). Thus, Maekawa solved the naming problem, but created confusing
nomenclature:

1) In China, tama no kanzashi ( = 玉簪) applies to the Chinese typical species,

which has shorter petioles, the leaves densely arranged, forming round clusters, the
leaf-blades being roundish, the narrow perianth tube (TA) is broader, the lobes of the
corolla are shorter and broader, and the ovaries fertile.

2) In Japan, tama no kanzashi ( = 玉簪) applies to the Japanese form, namely

H. plantaginea var. japonica (= タマノカンザシ = tama no kanzashi). This taxon is

longer stalked and the mound loosely arranged, the leaf-blades are oblong or elliptic-
ovate and not rounded and the lobes of the corolla and the narrow perianth tubes
(TA) are more narrow, and the plant is part-sterile (see herbarium specimens, page 3
─ see also the separate treatment of H. plantaginea var. japonica in this Species
Update). Flora of China (2004) does not recognize a Japanese form (= H.
plantaginea var. japonica. It is important to mention that in Japan the typical

Chinese H. plantaginea is called

マ ル バ タ マ ノ カ ン ザ シ (=

maruba tamano kanzashi).
According to ICBN rules, if H.
plantaginea var. japonica is
recognized, the full name of the
typical species will be H.
plantaginea var. plantaginea. A
large number of synonyms are on
record for this taxon. The specific
epithet plantaginea (meaning,
“resembling plantain”) likens this
species with cosmopolitan weeds
in the family Plantaginaceae.
Obviously not a weed, this species
has the most appealing and
fragrant flowers of any Hosta
species.

2010-02-05 ◄ H. plantaginea (cult.)

Flower Details of Typical Species
Court.: © kriss re kr ─ nature
Korea

-5-

H. plantaginea var. plantaginea (cult.)

A Group of Inflorescences in Full Bloom (Showing the Typical Horizontal Habit)
Photo Court.: ©baegdu.net Korea

Habitat and Biology: The introduction of H. plantaginea into Europe is well

documented (see pages 2 and 3). Unlike its European history and distribution, the
evolutionary habitat of this taxon in China is subject to supposition. Judging by
climate and length of season required for successful propagation among wild
populations (warm temperate and subtropical areas), some of the provinces named
below may not be suitable for propagation of endemic populations. Obviously,
cultivated specimens can be grown outside the required ecology. H. plantaginea
most likely evolved in the temperate and subtropical lower elevations of southern
China. Chinese folklore has a legend that a goddess dropped her hairpin from
which this beautiful hosta grew. While not science, it does confirm the presence of
and the love for this hosta among Chinese people. Documentation of use as a popular
garden plant, as food, as a medicinal herb and as a source for aromatics goes back
centuries to the Han Dynasty (漢朝; 206 BC–220 AD). Luo Yin (Zhaojian ─ 昭谏),
a Chinese statesman and poet near the end of the Tang Dynasty (618-904 AD) is

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mentioned in connection
with the use of this species.
While all of this is valuable
circumstantial and anecdotal
historical information, sci-
entific data relating to pre-
cise locations of natural
populations is unavailable.
Maekawa (1940), who re-
searched this subject with A.
Kikuchi, noted (page 346):
“Specimina spontanea nostris
ignota,” i.e., spontaneous
(wild) specimens [of this
species] are not known [to
us]. It is obvious that re-
search specimens used by
European and Japanese, as
well as Korean taxonomists
were from cultivated stock.
Flora of China (Vol. 24,
2000) states: “Forests,
grassy slopes, rocky places;
near sea level to 2200 m.

Anhui (安徽); Fujian (福建); H. plantaginea (European Form = ‘Grandiflora’?)
Guangdong ( 广 东 ; eastern
Inflorescences w/ Typical Horizontal Habit of
Guang); Guangxi (广西; wes- Flowers
tern Guang); Hebei (河)(?);
Hubei (湖北; north of lake); Institut für Pflanzenbiologie
Hunan (湖南; south of lake); © Court.: Technische Universität Braunschweig

Jiangsu (江苏); Liaoning (遼

寧)(?); Shaanxi (陕西(?); Qin Ling Mountains; Sichuan (四川); Yunnan (云南)

(?); Zhejiang (浙江)(?). Widely cultivated as an ornamental.” The question marks are

inserted by Flora of China editors signifying that spontaneous populations in the
listed provinces have not been confirmed. The evolutionary habitat mirrors plant
physiological-ecological requirements for successful seed production and this points to
the southeastern and southern provinces of China. Unconfirmed reports of

spontaneous populations in Zhejing province (浙江) would place such populations

further south than any other taxon in the genus Hosta. This taxon’s cultural

requirements tend to confirm such warm habitat. This also points to the postulation
that the highest habitat elevation listed in Flora of China (Vol. 24, 2000 = 2200 m

[7218 ft.]) may be too high an altitude for a suitable habitat. No spontaneous

populations are reported in Korea (KPNI - 국가표준식물목록 2004), but it is also

widely cultivated there. Specimens cultivated in North America are distributed over
a wide area, mostly in gardens. It is well known and documented (Schmid 1991; also

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in Hosta Journal; AHS) that in cultivation this species produces seed abundantly
during prolonged hot summers in southeastern North America. In the colder, more
northern areas, blooming is frequently aborted and the season is not long enough to
allow the production of viable seed. MOBOT Specimen Data Base shows 2 cultivated
North American specimens. USGA also lists cultivated specimen information. In
European gardens, this species has been cultivated since the 1780s and in warmer
regions, as for example along the northern Mediterranean coast, where the species
flourishes and produces abundant, viable seed (Schmid 1991).

Plant Morphology: Macromorphological, H. plantaginea is unique. As

shown on page 2, it develops the largest flowers in the genus Hosta. It is the only

species that is noticeably fragrant and it is night blooming. The best time to observe

the large flowers and notice the

fragrance is during evening

hours.

Conversion in Floral Archi-

tecture: A unique feature of H.

plantaginea is its unstable floral

architecture. Hosta flowers are

trimerous and consist of four

whorls of organs: 3 sepals, 3

petals, 6 stamens with anthers

and a pistil with a tripartite

stigma. Due to evolutionary

developments, in all hostas one

“conversion” has already taken

place. Its flowers have an outer

whorl (whorl 1) of 3 sepals,

followed by 3 petals (whorl 2).

Both the sepals and the petals

are alike and white or colored

and form the perianth so are

called tepals. This floral ar-

chitecture is typical in Hosta.

Floral identity (homeotic) genes

have been studied in other of H. plantaginea (Cult.)
Liliaceae (Hostaceae) and these
studies developed the so-called Early Conversion to 3 Petaloid Outer Stamens
ABC model. Explaining briefly, (This Is Not H. ‘Aphrodite’ but an Intermediate)
in the ABC model of floral organ
Hosta Hill R.G. © W.G. Schmid 1991.08.19

identity (Meyerowitz 1994), an A

function is responsible for the formation of sepals, the A+B for petals, the B+C for

stamen and the C-function for carpel formation. Further, Chaoing He, et. al. (2004)

report identification of homeotic floral mutants, which result in phenotypes showing

homeotic conversion of stamens into petals (▲see photo this page and page 9). Such

mutations affect organ identity of two adjacent whorls. This conversion is frequent

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Ovary and stamens

▼▼ ▼

44

◄ 18 Tepals

2010-02-05 ◄ Normal and petaloid stamens

H. ‘?[see page 10]’ © 2009 John Smead

-9-

in H. plantaginea, but rarely seen in other hosta species. Although not investigated
directly in Hosta, development of petaloid stamens in H. plantaginea are thus also a
result of a homeotic mutation as determined in other taxa in the Liliaceae, including
the true lilies. According to recent research, ABC-function genes encoding protein
transcription factors can form as dimers and/or tetramers of different composition
(Chaoing He 2004). This is based on a model called Floral Quartets (G. Theißen 2001),
which is also involved in the establishment of floral organ identity. Thus, several
factors are involved in the conversion of stamens or sepals to petals and the
production of multiple sets of tepals. In H. plantaginea, one of these conversions has
been described and named by Maekawa (1940; page 347) as H. plantaginea f.
Aphrodite (now considered a cultivar as H. ‘Aphrodite’; as reduced by W.G. Schmid

1991; page 85). It must be
stressed that the name H.
‘Aphrodite’ is applicable only
to Maekawa’s type with 3
exterior petaloid ►►►►►
►►►(continued page 11) ▼

◄ H. ‘???’

© Photo: J. Goodger/HL
◄◄◄ This form sold as H.
‘Aphrodite’ (which it is not!)

shows inner stamens with
anthers; shown here is not a
full conversion as described

by Maekawa but it is very
close to his 1940 description,
in which he did NOT specify
the number of tepals, hence

they are assumed to be a
normal number of 6 tepals.

◄ H. ‘Venus’

(Walters Gardens, Inc. and
C. Falstad 1993)

© Photo: K. Sisson/HL
◄◄◄ This cultivar shows 3

inner, slightly petaloid
stamens with 3 anthers

clearly visible; some
specimens of this cultivar
have no anthers showing and
the number of tepals varies.

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▲ Close-Up of 3 anthers ◄◄
Petaloid stamens with anthers

▼ H. ‘Venus’ ▲

(Walters Gardens, Inc. and C. Falstad; AHS 1993 ▪ © Photo: Meum71)
Examples of incomplete conversion of inner petaloid stamens with anthers attached

This Specimen has 16 Tepals

▼ Two flowers with 12 tepals and 6 petiolate stamens ▼
© Photo: R. Davis/HL

▼►►►►►►
(continued from
page10) stamens
fully developed
and the 3 inte-
rior stamens pe-
taloid, but shor-
ter and crispate
and 6 tepals not
18 as in photo on
Page 9). This
cultivar is unsta-
ble and exhibits
conversion of
inner stamens to
a full petaloid
form without
anthers, which

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has given rise to it being sold as “double-

flowered.” Many of the plants in the sold

as H. ‘Aphrodite’ have uncharacteristic

full double flowers and a number catalogs

and references (Zilis 2000, 2009)

characterize it as having full double

flowers, which is not the case. As

described by Maekawa (1940), it has 6

tepals, 3 large and 3 small petaloid

stamens. True doubling of the 6 tepals

would entail a doubling of the tepals to 12

to which petaloid or standard stamens will

be added. Some cultivars sold as H.

‘Aphrodite’ have 18! In 1993, C.H.

Falstad of Walters Gardens, Inc.

registered a cultivar that developed dur- H. plantaginea (cult.)

ing the production of H. plantaginea in Detail of Typical Floral Organs
tissue culture. This cultivar, named H. Anthers with Stamens and Pistil
‘Venus’, is described as “extra double
© W. George Schmid
huge flowers with 21 to 24 pure white

petals.” This indicates that: 1) the number of tepals is not constant, the maximum

being 24, the minimum being 12; and 2) the anthers are absent. The absence of

anthers is also reported in The Hosta Journal (K. Skrupky 1996). However, I have

examined many examples of H. ‘Venus’, which do have anthers carried on petaloid

stamens (page 10). This indicates that the organ conversion is not always complete. In

some conversions, petaloid stamens do have attached anthers that develop pollen.

Efforts to hybridize or self the multi-tepalous versions of H. plantaginea are rarely

successful (Zilis 2000, 2009).

Morphology: Plant size 60 cm dia. by 45 cm high (24 by 18 in.). Petiole 24 cm by

0.9 cm wide (10 by 0.35 in. wide) erect and spreading, unmarked glossy light green to
yellowish green (often darker green with the addition of nitrogen fertilizers). Leaf 16–
25 cm by 15–21 cm wide (6.5–10 by 6–8 in.), erect and in line with petiole, becoming
sub-pendulous, orbicular-cordate, nearly round, transition tight and contracted to
slightly open, small cuspidate, interior surface “flat,” but undulate, wavy in the
margin, not pruinose, surface dull, semi-glossy, below very glossy, light yellow green,
tip acuminate, projecting. Venation 8–9 (11), sunken above, very projected, smooth
below. Scape 60–80 cm long (24–32 in.), straight, erect, yellow-green, smooth round.
Fertile bracts, glossy green, ovate-lanceolate, grooved in the lower part, pointed; 2-
bracted, bracteole within external bract, opposite to lower fertile bracts, flat, large to
8 cm, fleshy and rigid, green or greenish white; lower, sterile bracts often larger and
foliaceous. Compact raceme 25 cm (10 in.), 10–15 flowers. NOTE: This is the only

species in Hosta with flowers degeneratively duplicate, with one-flowered, sessile
branches, appearing racemose, held erect in ±horizontal position on strong
pedicels, fragrant, very large, to 12.5 cm (5 in.) long and 7.5 cm (3 in.) broad, waxy,

shiny, lily-shaped, sharply recurving lobes, shiny white entirely, occasionally shades of

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white and pale cream, stamens not superior, parallel, joined to perianth tube;
conversion of stamens to petaloid shapes frequent; August, night-flowering. Anthers
yellow. Fertile, but requires correct timing of pollination due to night-blooming.
Capsules very large, 7cm (2.7 in) long by 7–8 mm (0.3 in.) wide, elongated, round,
with a pointed tip.

Karyotype-Chromosomes: Sporophytic Count = 60; 12 large and 48 small; diploid

(2n); (Mehra, P.N. & T.K. Pandita; Taxon 28:405 1979).

Pollen: Palynological data (M.G. Chung and S.B.

Jones; 1989; Pollen shape after Erdtman, 1966)

confirm correlations with the taxonomic treatment in

F. Maekawa (1940) and W.G. Schmid (1991) in

Subgenus Hosta (= formerly Subgenus Niobe). Pollen

morphology supports subgeneric and sectional

separation in Schmid (1991). The cited reference

indicates H. plantaginea has reticulate exine

ornamenta-tion with semitectate-columellate wall

struc-ture, while all the other species of Hosta have a

more or less continuous variation in the exine ▲ H. plantaginea var.
ornamentation, from rugulate to rugulate-baculate to japonica
rugulate-granulate exine ornamenta-tion with mostly
atectate wall architecture. In three specimens, no Pollen Type R (reticulate)
difference in the pollen type is found and all had Proximal Polar View of
pollen type R (reticulate). Note that two of the
Whole Grain

specimens had a similar pollen grain size, one had a

larger pollen grain size (sizes given in µm polar axis (P) × equatorial axis (E)):

Chung 227 (GA): = (P) 105.8 ± 7.4 to (E) 90.4 ± 5.4.

Bailey s.n. (BH): = (P) 105.3 ± 7.6 to (E) 96.4 ± 5.6.

Chung 234 (GA): = (P) 121.0 ± 3.9 to (E) 112.0 ± 4.3.

The Bailey specimen is determined to be H. plantaginea var. japonica and together

with Chung 227 gives a range size range for the variety. Chung 234 has been observed

by the author (at Dr. S.B. Jones Piccadilly Farm = source of the voucher) as being H.

plantaginea (forma typica). This specimen has a larger pollen grain, and the measured

value can be applied to this taxon.

H. plantaginea ►

Equatorial View
Showing Sulcus

SEM × 650
(M.G. Chung No. 1)

◄ H. plantaginea

Nonapertural Wall

TEM × 4000
(M.G. Chung No. 3)

2010-02-05 - 13 -

◄ H. plantaginea

Pollen Type R (reticulate)
Grain Surface Detail

SEM × 2000
(M.G. Chung)

DNA Banding: Cluster analysis of RAPD analysis

(Y. Yu, 2002; and Sauve, R.J., S. Zhou, Y. Yu, and
W.G. Schmid. 2005) has confirmed the genotypical
differentiation of H. plantaginea (F. Maekawa 1940;
in subgenus Niobe; W.G. Schmid in subgenus Hosta
1991). Comparison of the banding patterns and
cluster analysis supports the unique taxonomic
position of H. plantaginea as the type species of the
genus Hosta and its placement in the monotypic
subgenus Hosta. It further proves that H.
plantaginea is taxonomically differentiated from the
other species in the genus Hosta.

RAPD PCR Primer OPB-01: (5′-GTTTCGCTCC-
3′)

1 = H. plantaginea; 3 = H. venusta; 4 = H. minor;
5 = H. capitata; M = RAPD Markers.

Genome Size: DNA content (2C) in pg (10-12 gram) = 24.7 ± 0.26; Ratio PI/DAPI =

1.13. (Zonneveld, B.J.M. and F. Van Iren. 2001).

H. plantaginea (forma typica) 24.8 (= H. plantaginea)

H. plantaginea ‘Aphrodite’ 24.4 (= H. plantaginea conversion)

H. plantaginea ‘Richland Yae’ 24.4 (= H. plantaginea conversion)

H. plantaginea ‘Grandiflora’ 24.9 (= H. plantaginea var. japonica)

H. plantaginea ‘Yu Lei’ 24.9 (= H. plantaginea var. japonica)

These data show that no measurable difference exists in the DNA content between H.

plantaginea (forma typica) and H. plantaginea var. japonica (which see), indicating a

close relationship. Minor differentiation in morphological characters between these

taxa may be due to local adaptations over long periods and due to different habitat

ecologies. Two cultivars with floral conversion were tested and found to have the

same DNA (24.4 pg) content, which is less than the species. The latter may be due to

conversion factors, but that has not been comfirmed. It is confirmed that H.

2010-02-05 - 14 -

‘Aphrodite’ and H. ‘Richland Yae’ have the same origin. H. ‘Venus’ has not been
tested and macromorphological indicators show that it is H. plantaginea, which has
undergone multi-tepular conversion. Also not tested was H. plantaginea var.
stenantha, which is in this Species Update considered a selected cultivar form of H.
plantaginea var. japonica (which see), namely the cultivar H. plantaginea ‘Stenantha’
so not of taxonomic significance (Maekawa 1969; Schmid 1991).

Fertility in H. plantaginea. In 1939, Kono Yasui (保井コノ▪ the first woman

Doctor of Science in Japan) published an empirical study dealing with the (some-
times) problematic fertility experienced in H. plantaginea. A University of Tokyo
voucher of this species was 1) self-pollinated, and 2) pollinated with the pollen of H.
plantaginea var. japonica. The derived data determined the following:

1) Self-pollinated flowers produced pods with a good percentage of fertile seeds
and the calculated ratio of fertile to sterile seeds (aborted ovules) was 1:3.

2) The same plant when pollinated with the pollen of H. plantaginea var.
japonica produced a higher percentage of fertile seeds. Calculated ratio of fertile to
sterile seeds (aborted ovules) was 1:1.

Analyzing these data, Yasui determined that the taxon under examination is
a monogenic heterozygous plant of L and l; L symbolizes a dominant lethal gene
causing the partial sterility in this plant, and l is its allele, which is a recessive gene.
The combinations LL and Ll cause the abortion of the embryo and ll produces a
fertile embryo. The gene L does not affect any gametes. H. plantaginea var. japonica
is sometimes considered self-sterile but is actually a later season fertile plant that
requires a long season to produce viable seed. When H. plantaginea is pollinated
with the pollen of H. plantaginea var. japonica it produces viable seed because the
lethal L gene absent in the H. plantaginea var. japonica genetic makeup. Thus, the
ratio of viable to aborted ovules is 1:1, better than by self-pollinating the species.
Note that this was limited research and may not apply to all population samples of
H. plantaginea.

Taxonomic Type and Synonymy:

H. plantaginea (Lamarck) Ascherson (type of subgenus Hosta).

Botanische Zeitung, 21:53, ic. Pl. 24 1863.
Type: Ill. in Ascherson, Pl. 24, 1863 (syn.); Holotype in P; accepted taxon: GRIN

(USDA, ARS, National Genetic Resources Program); ITIS (Integrated
Taxonomic Information System: Taxon Serial No.: 42952); IPNI; MOBOT.

Botanical Synonyms:
Hemerocallis plantaginea Lamarck: Encyclopedie methodique Botanique, 3:103 1789.
Hemerocallis alba Andrews: Botanist. Repos. 3, tab. 194 1801.
Hemerocallis cordata Cavanilles: Descripcion de las plantas que Cavanilles, p. 124

1801.
Hemerocallis japonica in the sense of Redouté: Les Liliacées, 1:tab. 3 1802; in the sense

of Ker-Gawler apud Sims: Curtis's Botanical Magazine, 35, tab. 1433 1812
(with respect to the description and illustration; not Hemerocallis japonica as
originally described by Thunberg 1784 and 1794).

2010-02-05 - 15 -

Hemerocallis cordifolia hort. Salisbury: Transactions of the Linnean Society, London,
8:11 1807 (nom. nudum).

H. Iaponica (Thunberg; as incorrectly interpreted by Redouté and Ker-Gawler)
Trattinnick: Archiv der Gewächskunde, 1:55, ic. 89 1812 (excluding all the
synonyms).

Niobe cordifolia Salisbury: Transactions of the Horticultural Society, London, 1:335
1812 (nom. nudum and illegitimum).

Funkia subcordata Sprengel: Caroli Linnaei Systema Vegetabilium, Ed. 16, 2:40 1825
(nom. illegitimum and superfluum); and incl. Funkia var. grandiflora Siebold
ex Miquel: Verslag Mededelingen Akademie Wetenschappen, Amsterdam, Vol.
2, 3:304 1869 (excluding the synonyms by Banks and Houttuyn); Baker: J. of
the Linnean Society, London, 11:367 1870; Regel: Gartenflora, 25:162 1876;
deNoter: Rev. Horticol., Paris, N.S. 5, 77:390. ic. 163 1905.

Funkia alba (Andrews) Sweet: Hort. Brit., p. 409 1827.
Funkia sieboldiana bracteata Miquel: Annales Mus. Bot. Lugdano-Batavi, p. 152 1867

p.p. (nom. confusum).
Saussurea plantaginea (Lamarck) Kuntze (nom. nudum): Revisio Generum Plantarum

2:714 1891.
Funkia japonica (Thunberg; as incorrectly interpreted by Redouté and Ker-Gawler)

Mottet: Rev. Hortic., 69:115, ic. 37 1897; and incl. Funkia var. beta subcordata
(Sprengel) Lilja: Fl. Sveriges odl. vaexter, p. 49 1839; Druce: Second Suppl. to
Bot. Soc. and Exch. Club Rep. for 1916, p. 623 1917.
Hosta plantaginea f. grandiflora (Siebold and Zuccarini) Ascherson and Graebner:
Synopsis der Mitteleuropaeischen Flora, 3:53–55 1905 (not as pictured in
Lemaire 1846); Stearn: Gardener's Chronicle, 3 Ser., 90:48 1931.
Niobe plantaginea (Lamarck) Nash: Torreya, 11:3 1911.

H. plantaginea (Cult.)

Showing Typical Distended Flower Bud
© Hosta Hill R.G. ▪ Vouch.: Chung 234 W.G. Schmid; 1988.08.07

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Chinese, Japanese and Korean Synonyms:

玉簪 = ｷﾞｮｸｼﾝ = yu zan (Chinese)

棒玉簪 = ﾎﾞｳｷﾞｮｸｼﾝ = bang yu zan (Chinese Alias/Synonym)

玉春棒 = ｷﾞｮｸｼｭﾝﾎﾞｳ = yuchun bang (Chinese Alias/Synonym)

白鶴花 = ﾊｸｶｸｶ = bai he hua (Chinese Alias/Synonym)

マルバタマノカンザシ = maruba tama no kanzashi (Japanese; Kikuchi 1934)
옥잠화 = Ok-jam-hwa (Korean)
둥근옥잠화 = Dung-geun-ok-jam-hwa (Korean synonym)
비녀옥잠화 = Bi-nyeo-ok-jam-hwa (Korean synonym)

Horticultural Synonyms:

August Lily.

Corfu Lily Robinson 1883.

Fragrant Plantain Lily.

Funkia grandiflora hort.

Funkia liliiflora Foerster 1956 incorrect.

Funkia subcordata grandiflora hort.

Hosta ‘Grandiflora’ incorrect.

H. ‘Krossa No. K-2’ CL No. 318547

H. plantaginea subcordata grandiflora hort.

H. plantaginea ‘Long Tom’ hort. large flowers.

H. ‘Mack No. 14’.

Large White Plantain Lily.

Lilienfunkie (German). H. plantaginea (Cult.)
Paris Lily or Funkia.

Wegerichblätterige Funkie (German). Early Bud Stage

White Plantain Lily. © Hosta Hill R.G.

W.G. Schmid 1986.08.15

References:

Andrews, H.C., 1801. Rare and new Plants. Botanist. Repos. 3, tab. 194.

Ascherson, P.F.A., 1863. Botanische Zeitung, 21:53, Plate 24.

Ascherson, P.F.A., and K.O.R.P.P. Gräbner, 1905. Synopsis der mitteleuropäischen

flora, Leipzig, 3:53–55.

Bailey, L.H., 1903. Cyclopedia of American Horticulture. Ed. 2; 2:618-619

Baker, J.G., 1870. Funkia in: A revision of the genera and species of herbaceous

capsular gamophyllous Liliaceae. J. of the Linnean Society London, Bot. 11, 54–

Cavanilles, A.J., 1801. Descripcion de las plantas que D. Antonio Josef Cavanilles

demostró en las leciones públicas del año 1801. Madrid, p. 124.

Chaoing, H., T. Münster, and H. Saedler, 2004. On the origin of floral morphological

novelties. FEBS Letters 567:147–151 (Fed. of Euro. Biochemical Societies).

Chase, M.W. et al. 1995. Molecular phylogenetics of Lilianae. pp 109-137 in: Rudall

P.J., P.J. Crib, D.F. Cutler et C.J. Humphries (eds.). Monocotyledons:

systematics and evolution. Royal Botanic Garden, Kew.

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

and related genera. Bulletin of the Torrey Botanical Club, Vol. 116, 1:31–44.

2010-02-05 - 17 -

Conran, J.G. 1998 (in K. Kubitzki: Ed.):The Families and Genera of Vascular Plants,
Vol. 3.

Currie, H.E., 1988. Collected chemo-systematic data for a dissertation on
biosystematics of the genus Hosta. (Preliminary title: Biosystematics of the
Eastern Asian species of Hosta.). University of Georgia, Athens.

Davies, B., M. Egea-Cortines, E. de Andrade Silva, H. Saedler and H. Sommer 1996.
EMBO Journal 15:4330–4343.

Druce, G.C. 1917. Second Suppl. to Bot. Soc. and Exch. Club Rep. for 1916, p. 623.
Erdtman, G., 1966. Pollen morphology and plant taxonomy. Angiosperms. Hafner

Publishing Company, New York.

Flora of China 2000; Vol. 24:204, 205 (中國植物誌 – Chinese Edition)

Franchet, A. and P.A.L. Savatier, 1876. Enumeratio Plantarum in Japonia Sponte
Crescentium Hucusque Rite Cognitarum, Paris, pp. 82, 529.

Hensen, K. J. W., 1963. Identification of the hostas (“funkias”) introduced and
cultivated by von Siebold, Mededelingen van de Landbouwhogeschool te
Wageningen, Vol. 63, (6), pp. 6, 7, 11.

Iinuma, Y. (飯沼慾斎), (1856, 1874) 1910, Somoku Zusetsu (草木図説), An iconography

of plants, indigenous to, cultivated in or introduced into Nippon, revised by T.
Makino, Herbaceous Plants 6, pp. 462–470, pl. 19–27. Tokyo.
International Code for Botanical Nomenclature = ICBN, adopted by the International
Botanical Congress, Vienna Code 2005 (and earlier editions).
International Code of Nomenclature for Cultivated Plants = ICNCP, adopted by the
International Union of Biological Science, IUBS Division of Botany, 2004.

Korean Plant Name Index (국가표준식물목록 - Korean Edition 2004).

Kikuchi, A., 1934. Giboshi. 実用的園芸 (Practical Gardening), Vol. 17, 4:263–272.

Kuntze, C. E. O., 1891. Revisio generum plantarum 2:714.
Lamarck, J.B.P.A. de Monet de, 1789. Encyclopédie méthodique Botanique, Paris,

3:103.
Licht, J. L., 1989. Rediscovering an old H. plantaginea—Double the pleasure, twice

the fun. The Hosta Journal, Vol. 20, No. 2:11–12.
Lilja, N., 1839. Funkia var. beta subcordata. Fl. Sveriges odl. vaexter, p. 49.
Maekawa, F., 1937. Divisiones et plantae novae generis hostae (1). J. Japanese Botany,

13, No. 12:893–905.
Maekawa, F., 1940. J. of the Fac. of Science, Sect. 3 Botany, Vol. 5:342-348, ic. 6-12.
Maekawa, F., 1969. Hosta Trattinnick. In New Encyclopedia of Horticulture,

Seibundoshinkosha, Tokyo, Vol. 3:1107.
Matsumura, J., 1905. Index plantarum japonicarum, Tokyo, 2:200 (ex Miquel)
Mehra, P.N. & T.K. Pandita; Taxon 28:405 1979.
Meyerowitz, E.M. 1994. Commentary: Flower development and evolution: New

answers and new questions. Proc. Nati. Acad. Sci. USA; Vol. 91, pp. 5735-
5737, June 1994.
Miquel, F.A.W., 1867. Funkia sieboldiana bracteata. Annales Mus. Bot. Lugdano-
Batavi, p. 152 1867.
Moldenke, H.N., 1936. A monograph of the genus Cornutia. In Fedde Repertorium
novarum specierum regni vegetabilis, Berlin-Dahlem, 40:196–199.
Mottet, S.J., 1897. Funkia japonica. Rev. Hortic., 69:115, ic. 37.

2010-02-05 - 18 -

Nash, G.V., 1911. Niobe plantaginea. Torreya, 11:3.
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Company, Boston, p. 170.

Ranzan, Ono (小野蘭山) 1802/1803. Botanical classification (本草綱目啓蒙).

Ranzan, Ono (小野蘭山) 1847 (by Eds.) Revised and amplified edition (Jûtei Honzô-

Kômoku Keimô) of Botanical classification (本草綱目啓蒙).

Redouté, P.J. 1802. Les Liliacées, 1: tab. 3; in the sense of Ker-Gawler apud Sims:
Curtis's Botanical Magazine, 35, tab. 1433 1812 (with respect to the description
and illustration; not Hemerocallis japonica as originally described by Thunberg
1784 and 1794).

Regel, E. A. von, 1876. Die Funkia-Arten der Gärten und deren Formen. Gartenflora,
25:161–163.

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introduced since the death of Mr. Philip Miller. Transactions of the
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zeigen, 18.

H. plantaginea in
Cultivation:

H. plantaginea was the first hosta

grown in Europe (A complete H. plantaginea (Cult.)
nomenclatural history is presented
on page 1). Its distribution world- Typical Distended Bud Stage; Developing Buds
wide has revealed its preference
Court.: © Quingdao Nurs. 山東

for warmer and wet climatic 青岛平度苗圃

areas. Mirroring its (assumed!)

native habitat in China, this

species flowers in late summer

when the temperatures are high

and rainfall is abundant. At Hosta H. plantaginea (Cult.)
Hill in USDA Zone 7a; elevation
1188 feet (361m) AMSL; GPS Typical Distended Bud Stage; Developing Buds
coordinates 48.2083 W; 33.8500 N, Court.: © Quingdao Nurs. 山東
flowering time extends from mid-
青岛平度苗圃

August until mid-September,

depending on temperature and

precipitation. Given ideal conditions, H. plantaginea has the best flowers in the genus

Hosta. They are more than twice as large as those in other species, with a pure white

color, heavy substance and a waxy surface. This secies is the only night-bloomer with

flowers opening late in the afternoon. Their fragrance is most noticeable during

evening hours. The light green, shiny leaves provide a beautiful foil. Planting near

2010-02-05 - 20 -

garden seating or walks will

take full advantage this

species unique look and

lovely fragrance. Considered

by many a medium-sized

hosta it will grow into a large

clump in time. H. plantaginea

is parent to a number of

conversions and floral muta-

tions, which result in petaloid

stamens and multiple sets of

tepals (see pages 8-10). This

species requires a long season

with adequate rain, between

1.5 in/week (about 38

H. plantaginea (Cult.) mm/week) before and during

Mature Leaf Mound (Label = 3 × 1 in. (8 × 2.5 cm)) the bloom period. Extended,
© Hosta Hill R.G. W.G. Schmid 1986. warm summers are required
for productive set seed. It is

important to note that this

taxon is not the same one brought by von Siebold from Japan in the early 1840s,

which was H. plantaginea var. japonica with longer, narrower leaves and nar-rower

tepals (see page 3 and page 12 photo; this taxon is detailed in a separate section of

Species Update). Siebold's plant is most likely another morph of the species. While

most of the conversions result in enlarged and/or additional organs, the reverse is

possible.

H. plantaginea (Cult.)

Mature Leaf Mound (detail of sunken leaf veins) (margins damaged by drought)
© Hosta Hill R.G. W.G. Schmid 1985.08.20

2010-02-05 - 21 -

◄H.plantaginea

Note: Lack of water
during dry summer
periods, as seen here

in leaf margins
causes flowers to
diminish and occa-
sionally drop off.

Supplemental
watering is essential
during dry periods.

© W.G. Schmid
Hosta Hill 1985.08

Siebold’s “Japanese form” named Funkia grandiflora has been cultivated in Japan

for a very long time. First mentioned by the Japanese scholar Ranzan Ono (小野蘭山)

in Botanical classification ( 本 草 綱 目 啓 蒙 ; 1802/1803), it is now considered H.

plantaginea var. japonica. Notwithstanding, a form similar to that of von Siebold’s

Funkia grandiflora is still found in gardens under the name H. plantaginea ‘Grandi-

flora’. This is rather confusing,

because the typical species is also

sold under the name “Grandiflora”

in Europe. In 1940, Maekawa con-

firmed that the native Chinese

species with rounded leaves and

larger flowers was introduced into

Japan from Europe relatively

recently, with no exact date given.

Judging by the publication of the

Japanese names for H. plantaginea

var. japonica (タマノカンザシ =

tama no kanzashi (fide Ranzan Ono

ex Maek. 1940)) and H. plantaginea

f. stenantha (キダザキタマノカンザ

シ = kudazaki tama no kanzashi ex

Maekawa 1940) to differentiate

Japanese forms from the classic

Chinese species, the typical species

H. plantaginea may not have seen

widespread horticultural use in H. plantaginea var. japonica
Japan until the 1930s. Now, it is the
predominant form cultivated the Funkia grandiflora
world over, and has replaced H. Offered as H. plantaginea ‘Grandiflora’
plantaginea var. japonica in Japan- Note the Narrower Tepals and Slender Buds
ese public and private gardens.
© H. Philips/HL 2003

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Sports of H. plantaginea: mention a tepal number, which must be
taken as 3+3 = 6 tepals. This cultivar is
H. plantaginea has unstable occasionally seen with anthers attached
floral architecture as ex- to petiolate stamens due to incomplete
plained on pages 8-11. What conversion. H. ‘Venus’ is registered as
might be called the “basic” having “21 to 24” tepals, but due to
conversion (mutation) has incomplete conversion can be seen with
been described and named by as few as 12 tepals, and 3 large exterior
Maekawa (1940) and is now petiolate stamens and 3 inner petiolate
known as H. ‘Aphrodite’. He stamens with or without anthers. None of
describes H. ‘Aphrodite’ with the intermediate forms have been
3 large outer petiolate stamens separated out and named, which may be
and 3 inner short and crispate acceptable since only close examination
petiolate stamens without an- reveals the exact number of tepals and
thers as ►►► shown in the the overall multi-tepalous appearance is
photo (© T. Nakayama/HL) to not affected. Flowering time for the
the right. Maekawa does not conversion and TC mutants of H.
plantaginea remains the same as for the
H. ‘Heaven Scent’ typical species. Variegated sports of H.
plantaginea were unknown before tissue
© Walters Gardens, Inc. 1988 culture propagation and it can be stated
Hosta plantaginea TC Sport that its proclivity to produce variegated
sports is very low. Zilis (2000) reports
H. ‘Heaven Scent’ that “after propagating Hosta plantaginea
for several years in tissue culture, without
© Walters Gardens, Inc. 1988 seeing even a streak in a leaf, several
Hosta plantaginea TC Sport sports developed in my tissue culture
laboratory.” Zilis (Q&Z Nursery, Inc.)
introduced H. ‘Ming Treasure’ and H.
‘Fragrant Flame’ and C. H. Falstad (of

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◄◄◄ Typical Flower with 6
Tepals, 6 Stamens with Anthers,

and 1 Style with Stigma
(© J. Gamradt/HL)

◄▼ H. ‘Royal Standard’

Flowers Showing Early Stage
Conversion (Mutation) of

Stamens and Pistil with Outer
Stamens Showing Petaloid

Morphology © W.G. Schmid
▼▼▼ Hosta Hill R.G. 1985.08

(Walters Gardens, Inc.)
also introduced two of
tissue culture sports, H.
‘White Shoulders’ and H.
‘Heaven Scent’ (these
were awarded Plant
Patents No. 8015 and
8016, [with different
names]). H. ‘Chelsea Ore’
(Chelsea Physic Garden
1989) is often considered
a sport of H. plantaginea
‘Grandiflora’ (Grenfell
1996; p. 133), but it is not
fragrant and has laven-
der flowers. This points
to a hybridized origin
with the pod parent not
being H. plantaginea
(forma typica) but another cultivar. Aside from reported genetic and mutant
conversions of the flower architecture (see pages 8-12), sports of H. plantaginea retain
other dominant genetic characters. It is significant that the AHS Hosta Registry of the
International Registrar Kevin P. Walek does not show any direct sports of H.
plantaginea. All of the cultivars registered are sports of hybrids (!) of H. plantaginea.
The genetic predisposition for conversion of staminar tissue and multiplication of
tepals is occasionally transferred to hybrids with H. plantaginea as a pod parent, as
expected. H. ‘Royal Super’ (C.H. Falstad) is such a cultivar. It is described as a
“double-flowering,” variegated sport of H. ‘Royal Standard’ (♀ H. plantaginea × ♂
H. sieboldiana by J. Grulleman-Wayside/AHS 1986 (PP No. 2467). The latter is an
extremely popular and garden worthy cultivar that has been in cultivation for over
half a century. H. ‘Royal Standard’ is admired and found in gardens the world over.

2010-02-05 - 24 -

Propagation in Japan: The occurrence of “double-flowered” forms of H.

plantaginea has been known in China for a long time. In 1940, Maekawa was able to
obtain a specimen and named it H. plantaginea f. Aphrodite. Reports indicate that the
specimen was lost during WW II and it was several years after the end of the war
before this popular hosta again came to Japan. The Japanese love flowers so the
beauty and fragrance of the H. ‘Aphrodite’ flowers were an immediate hit with
gardeners and flower arrangers. Only a few plants were available and so rare and
and much sought after. The superiority of the Chinese H. ‘Aphrodite’ over the
Japanese H. plantaginea var. japonica was obvious. Recently, several news articles
(Shimbun 2005; Net Nihon Kai 2006) reported that in Tottori prefecture (鳥取県) of
the Chūgoku region (中国地方), rice farmers have changed rice crops to propagating
H. ‘Aphrodite’. The region’s climate is favorable and the farmers have found that
they can increase their farm income. Propagation methods, like TC are not available
to farmers so they use rhizome division.

Hybrid Progeny of H. plantaginea: The clone of H. plantaginea now cultivated

the world over, has been propagated vegetatively (by division and later tissue culture).
Its natural origin is unknown. Its resistance to successful hybridization is well known.
Although seed pods do form when this species is open pollinated or purposely
crossed with another species or cultivar, a substantial percentage of the seeds is not
viable. Notwithstanding, hybridizers have been able to successfully cross this species
and a number of hybrids are well known and are widely cultivated. Interspecific
crosses between other species of Hosta and H. plantaginea are difficult, but have been
successfully completed. One example is H. ‘Honey Bells’, which is a hybrid between
H. plantaginea (♀) and H. sieboldii (Schmid 1991). Chung and Jones (1989) pointed
out that the exine of this cultivar reveals a semi-tectate, columellate-like structure,
which mirrors that of H. plantaginea. Hybrids between H. plantaginea as pod parent
and other Hosta species show that the pod parents genes express dominance in
morphological, chemical, fragrance, and palynological traits. A list of H. plantaginea
hybrids is provided in the following, but note, that only direct species progeny with H.
plantaginea either as pod parent or pollen parent is included. If a H. plantaginea
hybrid is involved in a sport (H. plantaginea is part of the sport indirectly), or the
cultivar is a sport of a H. plantaginea hybrid sport, or for any other cultivars, in which
H. plantaginea is involved in a secondary role, consult the web site of the International
Registrar for the genus Hosta, Kevin P. Walek, at HostaRegistrar.org. A parent
search on this website will list all registered cultivars with H. plantaginea association.
Search HostaLibrary.org site for additional pictures.

The following abbreviations are used in the lists below on age 26:

♀ = the species as a pod parent = List 1

♂ = the species as a pollen parent = List 2

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List 1: Cultivars with H. plantaginea ♀ as a pod parent:

H. ‘Honeybells’ = ♀ H. plantaginea × ♂ H. sieboldii by A. Cumming/AHS 1986
H. ‘Iron Gate Supreme’ = ♀ H. plantaginea × ♂ H. ‘Tokudama Aureonebulosa’ by

V. Sellers 1980
H. ‘Royal Standard’ = ♀ H. plantaginea × ♂ H. sieboldiana by J. Grulleman-

Wayside/AHS 1986 (PP No. 2467)
H. ‘Willamette’ = ♀ H. plantaginea × ♂ H. ‘Royal Standard by P.& J. Ruh 2003

List 2: Cultivars with H. plantaginea ♂ as a pollen parent:

H. ‘Austin Dickinson’ = ♀ H. ‘Resonance’ × ♂ H. plantaginea by W. & E. Lachman
1992

H. ‘Buckwheat Honey’ = ♀ H. ‘Ginko Craig’ × ♂ H. plantaginea by R. H. Benedict
1984

H. ‘Curly Top’ = ♀ H. ‘Ginko Craig’ × ♂ H. plantaginea by R. H. Benedict 1985
H. ‘Emily Dickinson’ = ♀ H. ‘Neat Splash’ hybrid × ♂ H. plantaginea by W. & E.

Lachman 1987
H. ‘Flower Power’ = ♀ H. nigrescens × ♂ H. plantaginea by K. Vaughn 1987
H. ‘Forever Green’ = ♀ H. rupifraga × ♂ H. plantaginea by J. Dishon 1994
H. ‘Green Marmalade’ = ♀ H. ‘Neat Splash’ × ♂ H. plantaginea by C. Owens 1987
H. ‘Mistress Mabel ’ = ♀ (H. ‘Beatrice F6’ × H. ‘Blue Moon’) × ♂ H. plantaginea by

W. & E. Lachman 1995
H. ‘Seventh Heaven’ = ♀ H. ‘Kevin Vaughn’ × ♂ H. plantaginea by J. Kulpa 1994
H. ‘Slim Polly’ = ♀ H. sieboldii × ♂ H. plantaginea by F. Williams 1986
H. ‘Snow Flakes’ = ♀ H. sieboldii ‘Alba’ × ♂ H. plantaginea by F. Williams/AHS

1986
H. ‘Summer Fragrance ’ = ♀ H. ‘Vaughn 73-2’ × ♂ H. plantaginea by K. Vaughn

1983
H. ‘Sweet Marjorie’ = ♀ H. ‘Ginko Craig’ × ♂ H. plantaginea by R. H. Benedict

1983
H. ‘Sweet Susan’ = ♀ H. sieboldii × ♂ H. plantaginea by F. Williams & C. Williams

1986
H. ‘Warwick Essence’ = ♀ H. ‘Northern Halo’ × ♂ H. plantaginea by L. Jones 1993

NOTE: The above lists do not include the many Japanese cultivar names
associated with H. plantaginea. Most of these names are described earlier
under the section H. plantaginea in Cultivation. There are a number of
Chinese and Japanese cultivars and some have been mentioned earlier under
their respective names. The main goal of Japanese cultivation is the flower,
which is used for flower arranging and show displays. These intermediate
forms are not generally available so are not detailed nor mentioned.

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H. plantaginea f. Aphrodite Maek. = H. ‘Aphrodite (cultivated)

Garden Setting in Gosen-shi (五泉市), Niigata-ken (新潟県)
© Photo: Toyozo Nakayama

H. plantaginea f. aphrodite Maekawa 1940

J. of the Faculty of Science, Imperial University Tokyo, Sect. 3 Botany, Vol. 5:343,
344, 347, 348, ic. 11 1940.

H. plantaginea ‘Aphrodite’ Schmid 1991

The genus Hosta: Gibōshi Zoku (ギボウシ属); Timber Press; Portland. P. 85.
雙花玉簪 = ching ban yuzan (Chinese = H. ‘Chingbanyuzan’)

八重 咲きマルバタマノカンザシ = yae no maruba tama no kanzashi (Japanese)

History, Nomenclature, and Synonymy: H. plantaginea f. aphrodite was

described as a taxon by F. Maekawa (1940). W.G. Schmid (1991) reduced this taxon
to cultivar status, because 1) it developed in cultivation among cultivated material,
and 2) it can not be propagated by way of seed due to its infertility. The following
additional historical and nomenclatural information is included here applying
specifically to this cultivar, for which the name H. ‘Aphrodite’ is in garden use: In
1940, F. Maekawa established and described a new forma under the species H.
plantaginea. He named it H. plantaginea f. Aphrodite. The new taxon was based on a
living, cultivated plant received from China, Hebei province (河北; Hébĕi) growing
as No. 320 in Hortus Kikuti. Coming from a northern province in China, it was

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from cultivated stock. Maekawa

assigned a Japanese name com-

bination: 八 重 and 咲 き ; the

combination meaning “double

flower” (八重 咲きマルバタマ

ノカンザシ; transliterated and

meaning Yae-no-maruba tama-

no-kanzashi; the transliteration

is seen simplified to Yae-tama-

no-kanzashi). In China it is

referred to as 雙花玉簪 = ching

ban yuzan and this trans-

literation has found its way into

horticultural use as H. ‘Ching-

banyuzan’, which should be

considered a synonym. As a

taxon, it has also been named in

China as H. plantaginea var.

plena by Yan Liang Fei (1983;

cfr. Licht 1989) but Maekawa's

name H. plantaginea var. H. ‘Aphrodite’
aphrodite has priority so the
former name is a nomen super- 3 Open Flowers
fluum. Following this, in North © W.G. Schmid Hosta Hill R.G. 1994.08

America it was known as H. plantaginea ‘Flora Plena’ or H. plantaginea ‘Plena’ but

these names are per the ICNCP invalid because they are based on invalid names and

this taxon should be called H. ‘Aphrodite’, per Maekawa (1940). The exact date of the

discovery of multi-tepal (for example double-flowered) and other forms like H.

‘Aphrodite” is unknown but may have occurred in China among cultivated plants

many years ago and it is known that H. ‘Aphrodite’ came from China in the 1930s

(Maekawa 1940). His specimens were lost during WW II. Even now similar cultivars

come from China under different names, for example H. plantaginea ‘Yu Lei’, which

is also known as H. ‘White Fairy’. The name Yu Lei comes

from the Chinese 簪 (meaning “ornamental hairpin”) and

the Kanji 蕾 (equivalent to Chinese Pinyin lěi, meaning

bud) so the two combined stand for ◄◄◄ “ball bud.”

The large bud often takes on a ball shape, hence the name.

In this case, the Chinese name should be used because it

was established in China before the cultivar name H.

‘White Fairy’ appeared. Nowadays, conversion of stamens

to petals occurs even in gardens and is not a new

phenomenon and I have observed it at Hosta Hill R.G. on several occasions and many

inter-mediate flowering forms can be found.

Morphology: As described by Maekawa (1940; page 347), the name H. ‘Aphrodite’

applies only to a plant with 3 exterior petaloid stamens fully developed and the 3 inte-

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rior stamens petaloid, but shorter and crispate and six 6 normal tepals. Any con-
versions with more than 6 tepals are NOT H. ‘Aphrodite’ (see also pages 9-12).

Taxonomic Type and Synonymy:

H. plantaginea `Aphrodite' (a cultivar form as H. ‘Aphrodite’): The genus Hosta:
Gibōshi Zoku (ギボウシ属); Timber Press; Portland. P. 85.

Botanical Synonyms:
H. plantaginea f. aphrodite Maekawa: J. of the Faculty of Science, Imperial University

Tokyo, Section 3 Botany, Vol. 5:347 1940 (reduced to cultivar form).
H. plantaginea var. plena Fei: Huahui Ji Guanshang. Shouce Press, Taipei, 1983.

Chinese and Japanese Synonyms:
雙花玉簪 = ching ban yuzan (Chinese = H. ‘Chingbanyuzan’)
八重 咲きマルバタマノカンザシ = yae no maruba tama no kanzashi (Japanese)

Horticultural Synonyms:
Double-flowered White Plantain Lily.
H. plantaginea ‘Aphrodite’.
H. plantaginea ‘Flora Plena’ incorrect.
H. plantaginea ‘Plena’ incorrect.
H. plantaginea ‘Yu Lei’ (‘White Fairy’).

2010-02-05 ▲ Bracts with Bud Initial
Hosta Hill R.G.

© W.G. Schmid; 1983.06

◄▲H. ‘Honeybells’

◄ Leaf Mound with Ligularia
stenocephala ‘Weihenstephan’

Hosta Hill R.G.
©W.G. Schmid 1983.06

- 29 -

Some H. plantaginea Hybrids in Cultivation

Some of the oldest known hybrids of H. plantaginea are classics found in many
gardens. H. ‘Honeybells’ is often referred to as a “hosta weed” because it multiplies
fast and adapts to just about any gardening condition and setting. Another classic is
H. ‘Royal Standard’, with thicker leaves and also a fast growth rate. Its leaves have
more substance and are attractive. Both have fragrant flowers. H. ‘Honeybells’ has
lavender flowers and they are white in H. ‘Royal Standard’. Both make great
ground covers. They provide huge numbers of flowers in late summer for 3-4 weeks.

H. ‘Honeybells’ H. ‘Royal Standard’

Two classic H. plantaginea hybrids used as ground cover.

Left: H. ‘Honeybells’ with white-margined H. ‘Undulata Albomarginata’ used as

foundation planting at Hosta Hill R.G. (© W.G. Schmid; June 1984). Right: H.

‘Royal Standard’ used as ground cover and tree underplanting (© W.G. Schmid;

June 1987).

In both gardens, these ground covers become a floriferous, fragrant display in mid-
August and carry on for almost a month. Both hybrids are readily available and
inexpensive, which makes such displays possible to most gardeners. In the case of
Hosta Hill R.G., 6 plants were obtained in 1969 and were divided at frequent
intervals, resulting in hundreds of “free” plants by 1984.

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H. ‘Honeybells’

Elongationg Raceme with Bud Initials and Branching Scape
Hosta Hill R.G. 831008001; © W.G. Schmid; 1983.08.01

H. ‘Honeybells’

Mass Planting Showing Developing Racemes and Buds
Hosta Hill R.G. 831008001; © W.G. Schmid; 1983.08.01

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▲ H. ‘Royal Standard’

Developing Flower Bud and Raceme
Hosta Hill R.G. © W.G. Schmid; 1987.08

◄ H. ‘Honeybells’

Elongating Raceme with Opening Flowers

©W.G. Schmid 2006/2010: The text and illustrations are
copyrighted and are available for personal reference only.
The content may not be published in printed form without

the author’s written permission. Web quote reference:
W. George Schmid, HostaLibrary.org/species/.

Note:
Also refer to the section on

H. plantaginea var. japonica

which includes additional comparative
information

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