241
A eapp aisal o Thunbe g’s spo ed lily Lilium macula um
Thunb. and ecogni ion o he elegan lily Lilium × elegans
Thunb. (Liliaceae)
James A. Comp on1
1 Spilsbu y Fa m, Salisbu y, SP36RU, UK
Co esponding au ho : James A. Comp on ([email p o ec ed])
Copy igh : © James A. Comp on
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Thunbe g encoun e ed nine species o Lilium du ing his isi o Japan in 1776. O hese,
L. lanci olium Thunb., L. longi lo um Thunb., L. macula um Thunb., and L. speciosum
Thunb. a e all accep ed. He also desc ibed L. elegans Thunb., a lily ha is mo pholog-
ically simila o L. macula um. The long-dispu ed name L. ×elegans is con i med as
co ec ly belonging o he hyb id comp ising L. macula um and L. pensyl anicum Ke
Gawl. Thunbe g’s wo king me hodology, including ha o his illus a o s and eng a -
e s, is sc u inised conce ning he publica ion o his Japanese lilies. Fou lec o ypes and
a neo ype a e designa ed, a ull synonymy is lis ed o L. macula um and L. ×elegans,
and keys o he hyb id and i s pa en species, as well as o he na u al a ie ies wi hin
L. macula um, a e p o ided.
Key wo ds: Dejima, Japan, Lilium × elegans, L. macula um, nomencla u e, Thunbe g
In oduc ion
Six een ecognised species o he genus Lilium L. occu in Japan (Hayashi
2016: 111). Two o hese, Lilium macula um Thunb. and L. pensyl anicum Ke
Gawl., ha e sca e ed lea es and up igh , bowl- o cup-shaped lo al mo phol-
ogy wi h epals ha na ow o a claw-like base. Lilium macula um is endemic
o Japan, while L. pensyl anicum has a much wide dis ibu ion ac oss much
o empe a e o sub-a c ic eas e n Asia (Comp on and Sy in 2023). Recen
molecula da a ha e shown ha hei nea es ela i e is he Eu opean species
L. bulbi e um L. (Du e al. 2014: 254; Huang e al. 2018: 1253; Gi nish e al.
2020: 38; Wa anabe e al. 2021: 193). These da a ha e also shown ha he
wo Asian species and he single Eu opean species belong oge he wi h se -
en Asian species in L. sec . Sinoma agon H.F.Combe , and ha e also been
placed oge he in L. subsec . Dahu ica Ba ano a, classi ica ions based o ig-
inally on mo phology (Combe 1949: 101; Ba ano a 1988: 1327). Lilium sec .
Sinoma agon is now cha ac e ised as ha ing ad en i ious s em oo s abo e
he bulb; s ems mo e o less s ia ed; a ma ginal idge on ei he side o a
sinus on he adaxial su ace o he epals; and sho papillae p esen on he
s igmas (Wa anabe e al. 2021: 199, 201).
Academic edi o : Lo enzo Pe uzzi
Recei ed:
30 June 2025
Accep ed:
16 Sep embe 2025
Published:
9 Oc obe 2025
Ci a ion: Comp on JA (2025) A
eapp aisal o Thunbe g’s spo ed
lily Lilium macula um Thunb. and
ecogni ion o he elegan lily
Lilium × elegans Thunb. (Liliaceae).
Phy oKeys 263: 241–270. h ps://doi.
o g/10.3897/phy okeys.263.163611
Phy oKeys 263: 241–270 (2025)
DOI: 10.3897/phy okeys.263.163611
242
Phy oKeys 263: 241–270 (2025), DOI: 10.3897/phy okeys.263.163611
James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
Two No h Ame ican species, L. ca esbaei Wal e and L. philadelphicum L.,
sha e se e al simila lo al mo phologies wi h he Eu asian species, such cha -
ac e s leading bo anis s in he pas o classi y hem oge he unde wha John
Gilbe Bake e e ed o as G oup 3 “Isoli ion” (Bake 1871: 104; Wallace 1873:
63). Acco ding o molecula da a (Du e al. 2014: 254; Huang e al. 2018: 1253;
Gi nish e al. 2020: 38; Wa anabe e al. 2021: 193), hese wo No h Ame ican
species a e no closely ela ed o he Eu asian species and belong oge he in
a di e en sec ion o he genus, Lilium sec . Pseudoli ium (Endl.) Spach (Spach
1846: 277). Close examina ion o hese No h Ame ican species e eals ha
hey can also be dis inguished om he Eu asian species by hei glab ous
s ems, lea es, and lowe buds, and by hei epals, which na ow ab up ly o-
wa ds he base, o ming a linea u ow.
Lilium bulbi e um, he only Eu opean species in L. sec . Sinoma agon, is
cha ac e ised by ha ing small bulbils p oduced in he lea axils along he in lo-
escence axis, al hough bulbils may be lacking in a . c oceum (Chaix) Pe s.,
bu wi h e e e s ems wi h less pubescence on he in lo escence axis han
ha on i s Asian coun e pa s. This species, ypi ied on a specimen clea ly
ha ing bulbils LINN 420.2 (Pe uzzi and Ja is 2009: 1361), is undoub edly
closely ela ed o bo h L. macula um and L. pensyl anicum, sha ing simila
mo phologies and molecula da a (Du e al. 2014: 254; Huang e al. 2018:
1253; Gi nish e al. 2020: 38; Wa anabe e al. 2021: 193). I is, howe e , e-
s ic ed o Eu ope and is no he ocus o his pape .
The e is li le doub ha L. macula um and L. pensyl anicum closely e-
semble each o he mo phologically. Thei geog aphical dis ibu ional anges
only o e lap in he mos no he n ex emi y o Honshu Island, whe e L. pen-
syl anicum has i s mos sou he ly popula ions in Japan (Hayashi 2016: 111,
112; Wa anabe e al. 2024: 467). Lilium pensyl anicum has ibbed s ems wi h
loccose pubescence dis ibu ed on he uppe s em pa s, pa icula ly in he
lea axils and along he lowe buds and pedicels, and has hypogeal seed
ge mina ion. Lilium macula um has sca ious s ems wi h papillose hai s and
epigeal seed ge mina ion (Hayashi 1990: 9; 2016: 112; Wa anabe e al. 2024:
460). Lilium pensyl anicum occu s h oughou much o cen al and eas e n
Sibe ia, he Ko ean Peninsula, Manchu ian China, and Mongolia. I also oc-
cu s in Sakhalin and he sou he n Ku il Islands (Comp on and Sy in 2023). In
Japan, i is ound ac oss Hokkaido and in a ew places in he ex eme no h
o Honshu, on Shimoki a Peninsula and a ound Cape Tappi on he Tsuga u
Peninsula, bo h in Aomo i P e ec u e (Hayashi 2016: 112).
A ecen conse a ion p oposal by Y-D. Gao o conse e he name L. dau-
icum Ke Gawl., a name widely used in he pas , o e L. pensyl anicum (Gao
2021: 1139–1140), has been ejec ed (Applequis 2023: 917–918). This con-
clusion means ha L. pensyl anicum emains he co ec name o ha eas
Asian species. The ull ange o names associa ed wi h L. macula um is dis-
cussed below (see synonymy).
The eas Asian mo ning s a lily, L. concolo Salisb., which also belongs
in L. sec . Sinoma agon, was so named o i s lowe s wi h much na owe
epals which adia e in a s ella e manne (Hayashi 2016: 112). An a icle by
F i z Be ckmülle o Hambu g indica ed ha Thunbe g’s L. macula um (unde
he name L. hunbe gianum) was o hyb id o igin wi h L. pensyl anicum (as
L. dau icum) and L. concolo as he pa en species (Be ckmülle 1927: 216).
243
Phy oKeys 263: 241–270 (2025), DOI: 10.3897/phy okeys.263.163611
James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
This hyb id iden i y was hen aken up unde he name L. ×macula um by la e
wo ke s (e.g. S ea n 1948: 12; Woodcock and S ea n 1950: 263; Synge 1980:
64). The wide ange o a ia ion exhibi ed in ho icul u al selec ions o plan s
p e iously iden i ied as L. macula um is likely o ha e enhanced ha conclusion.
Cha ac e s such as obus ness o plan s a u e, inc eased numbe o lowe s,
a iable epal colou ing, and o namen a ion on he epals a e s ong indica o s
o po en ial hyb idi y. A e y simila lily o bo h L. macula um and L. pensyl an-
icum, desc ibed by Thunbe g as L. elegans Thunb. (Thunbe g 1811: 203), has
also been pos ula ed o be o hyb id o igin wi h L. pensyl anicum and L. macu-
la um as he pu a i e pa en species (e.g. Shimizu 1960: 105; 1969: 27; 1973:
119; Fu uya 2009: 85; Okubo 2014: 12). Recen esea ch based on mo phology
as well as DNA sequence da a om plas id and nuclea genes and gene space
egions has con i med he exis ence o his hyb id, al hough no hyb id o mula
was p o ided in ha wo k (Wa anabe e al. 2024: 457, 467). In his pape , he
hyb id s a us o bo h L. ×elegans and L. ×macula um is discussed. A c i ical
o e iew o Thunbe g’s wo king me hods, a e iew o his publica ions, and a
ho ough examina ion o his he ba ium specimens wi h espec o Japanese
Lilium names has been unde aken. An app aisal is made o all he subsequen
names associa ed wi h hese lilies, and hei s a uses a e assessed.
Thunbe g’s disco e y o he spo ed lily
Lilium macula um Thunb. was i s desc ibed by he Swede Ca l Pe e Thun-
be g (1743–1828) in he second olume o he B i ish jou nal T ansac ions
o he Linnean Socie y o London (Thunbe g 1794: 334). His desc ip ion
o he species was made wi hin he commen a y on his majo wo k, Flo a
Japonica, published a decade ea lie (Thunbe g 1784). Thunbe g, who was
one o Linnaeus’s s uden s, collec ed ma e ial o he lily in he summe o
1776 du ing he 15 mon hs which he spen in Japan. I was no possible a
ha ime o “ o eigne s” o a el anywhe e in Japan. The Chinese we e pe -
mi ed a small encla e ou side he ci y o Nagasaki, and he Du ch we e e-
s ic ed nea by o he small a i icial island o Dejima in Nagasaki Bay. This
island, which was o all in en s and pu poses a iny Du ch colony, was linked
by a gua ded b idge o he la ge sou he n Japanese island o Kyushu. The
Du ch we e he only Eu opean na ion pe mi ed by he Japanese o ade
wi h hem, bu only om his small island. Thunbe g’s a els while he was
in Japan, always unde a Japanese esco , we e consequen ly es ic ed o
he locali y a ound Dejima, he excep ion o him being he annual ho eis o
cou jou ney, an expedi ion unde aken by he Du ch in o de o pay homage
o Japan’s mili a y ule , he shōgun, in Edo [Tōkyō] on Honshu. This ule o
es ic ed mo emen came unde he sakoku o “locked in” policy, which ex-
cluded all o eigne s om se ing oo on Japanese soil wi hou pe mission
and was no e oked un il almos 100 yea s a e Thunbe g’s ime he e.
Al hough he was Swedish and had se e al eminen Swedish sponso s,
including Linnaeus, his jou ney o Japan was sanc ioned and sponso ed on
behal o he Du ch Eas India Company (VOC) by se e al in luen ial Ams e -
dam bu ghe s, including his con empo a y he alde man Jean [Johann] Deu z
(1743–1788), a di ec o o he Su inam Socie y and co esponden o Joseph
Banks; bu gemees e Egbe de V ij Temminck (1700–1785); and he ci y
244
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
councillo , di ec o o he Su inam Socie y, and bu gemees e Jan an de Poll
Pie e sz (1726–1781). Also much in ol ed in p omo ing Thunbe g’s employ-
men as a physician o he VOC was Linnaeus’s iend and colleague he Du ch
bo anis Johannes Bu man (1707–1780), p o esso a he Ams e dam A he-
naeum Illus e [la e Uni e si y o Ams e dam] (Skuncke 2014: 58). Thunbe g
was commissioned ini ially o isi he Cape in Sou h A ica, whe e, om 1772
o 1775, he lea n Du ch. His inal des ina ion, howe e , was Japan, whe e he
would be based as he physician in he Du ch ac o ij o ading pos on Dejima,
which Thunbe g soon ound ou was o become a i ual p ison.
The Japanese name o Lilium macula um is sukashi yu i スカシユリ, a ph ase
meaning “see- h ough lily”, a e e ence o he open spaces be ween he bases
o each epal. The species is es ic ed o Honshu, g owing on sandy banks and
ocky ledges om Tōhoku in he no h o he island sou hwa ds o he cen al
Chūbu egion (Hayashi 2016: 112). Thunbe g ne e isi ed he Tōhoku egion
as i was a oo dis an om he places he was pe mi ed o a el in he sou h-
e n pa s o Japan. He did, howe e , pass h ough eas e n coas al pa s o he
Chūbu egion in Honshu (see below).
The annual ho eis o cou jou ney mus ha e been a g ea escape om
Thunbe g’s con inemen . Each yea , he Du ch on Dejima had o pay homage
o he de ac o ule o Japan, he shōgun Tokugawa Ieha u, in a away Edo
on Honshu. This homage in ol ed a land and sea jou ney co e ing a dis ance
o 2,600 km, which, including he e u n jou ney, las ed o se e al mon hs. On
4 Ma ch 1776, he Du ch chie me chan on Dejima, A end Willem Fei h, ac-
companied by Thunbe g, as physician o he Du ch, and he sc ibe He man
Köhle , se ou on hei jou ney. Thei ou e ook hem along he Paci ic coas o
he Chūbu egion o Honshu, h ough pa s o mode n-day Shizuoka P e ec u e
whe e he lily s ill g ows (Kijewski 2015: 162, map ig. 2).
Thunbe g ecoun s his jou ney in he P e ace o Flo a Japonica (Thunbe g
1784: x iii):
“Annus 1776 mihi e Flo ae dilec issimae maxime a eba ; i e enim ad au-
lam se o suscep um, soli o diu ius p o aheba u , ob ad e sos in ma i en os
130 millia ibus p opius ad li o a con ec is. Hinc e p ae e iba , aes as e o
ad en aba , u in edi u plu imas inuene im plan as eximie lo en es.Dum in
Me opoli Iedo e saba , ab 18 Maii ad 13 Jun. in di e so io saepissime me
adiban duo medici Ka agawa Fosiu e Nakagawa Sunnan, in Mine alogia,
Zoologia, Bo anica e Medicina haud pa um e sa i [The yea 1776 was mos
a ou able o me and o my belo ed Flo a; o he cou jou ney, ha ing been
unde aken la e, ex ended longe han usual, owing o ad e se winds a sea,
we ound ou sel es o 130 miles close o he coas . F om he e, he sp ing
passed away, and he summe came on, so ha on my e u n, I ound many
plan s blooming exquisi ely. While I was in he me opolis o Yedo (Tokyo),
om he 18 h o May o he 13 h o June, wo doc o s, Ka agawa Fosiu and
Nakagawa Sunnan, who we e well e sed in mine alogy, zoology, bo any, and
medicine, isi ed me e y o en in my dwelling] (Thunbe g 1784: x iii).
His ansla ion o he Japanese names Ka agawa Fosiu and Nakagawa Sun-
nan was p obably in ended o e e o he physician and ansla o Ka su agawa
Hoshū (1751–1809) and he su geon Nakagawa Jun’an (1739–1786). As well
245
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
as wild plan s in Shizuoka, i is also likely ha Thunbe g saw lowe ing plan s o
L. macula um and L. elegans cul i a ed in he ga dens whe e e he had s opped
along he ou e du ing his e u n jou ney om Edo ia Miako [Kyō o] and Ōsaka
on Honshu on his way back o Dejima, Nagasaki, on Kyushu.
Thunbe g’s Flo a Japonica
Thunbe g’s i s con ibu ion o he classi ica ion o he lilies o Japan was in his
Flo a Japonica (Thunbe g 1784: 133–135). O he eigh lilies he had included
(Table 1), only one species, L. japonicum Thunb. ex Hou ., s ill has he co ec
name (Thunbe g 1784: 133). I was desc ibed in 1780 by he Du ch bo anis
Maa en Hou uyn om one o Thunbe g’s collec ions and published ou yea s
be o e Thunbe g’s own accoun (Hou uyn 1780: 245). P io o his a i al in Ja-
pan, Thunbe g’s knowledge o he genus Lilium in ha coun y was ine i ably
limi ed o accoun s ha had been p e iously p o ided by a hand ul o Eu opeans
who had been able o gain access o some knowledge abou Japanese plan s.
In his con ex , h ee o Thunbe g’s species names a e impo an in helping o
unde s and his concep o wha hese Japanese species may ha e ep esen -
ed o him: he Eu opean L. bulbi e um L. (Linnaeus 1753: 302) and wo No h
Ame ican species, L. canadense L. (Linnaeus 1753: 303) and L. philadelphicum
L. (Linnaeus 1762: 435). His ini ial species de e mina ions in Japan, he e o e,
we e in luenced by his unde s anding o species which had p e iously been de-
sc ibed om elsewhe e in he wo ld. Subsequen examina ion o his he ba ium
specimens, ca e ully sc u inised along wi h he anno a ions on his shee s, has
p o ed pi o al in co ec ly de e mining he iden i y o se e al o his axa.
When conside ing Thunbe g’s de e mina ions, i should also be bo ne in
mind ha a e he had le Japan on 3 Decembe 1776, he did no e u n o
his na i e Sweden o o e 3 yea s, ha ing spen ime a elling and collec -
ing in Ja a, S i Lanka, he Cape o Good Hope, and Sie a Leone. Mo eo e ,
he also spen 2 mon hs s udying in Ams e dam and London, inally eaching
Ys ad in sou he n Sweden on 14 Ma ch 1779 (Skuncke 2014: 186). A e his
a i al in Sweden, i was ano he 5 yea s be o e his Flo a Japonica was e en-
ually published in he Uppe Saxon ci y o Leipzig (Thunbe g 1784). O e ha
pe iod, some o his no es we e likely o ha e become con used, and his ini-
ial ecollec ions o he axa conce ned may ha e become less han accu a e.
Table 1. Ch onology o Thunbe g’s de e mina ion and naming o Japanese Lilium species.
Flo a Japonica 1784 Obse a ions 1794 Examen 1811 Cu en ly accep ed name and specimen in UPS
L. bulbi e um L. lanci olium L. lanci olium L. lanci olium V-008139
L. canadense – – L. leich linii subsp. maximowiczii
L. candidum L. longi lo um L. longi lo um L. longi lo um V-008140
L. japonicum a . α –L. japonicum L. japonicum V-008137
L. japonicum a . β – – L. japonicum
L. philadelphicum L. bulbi e um L. elegans L. ×elegans V-008129
L. pomponicum –L. pomponicum L. concolo V-008147
L. supe bum L. speciosum L. speciosum L. speciosum V-008149
–L. co di olium L. co di olium Ca dioc inum co da um V-008135
–L. macula um L. macula um L. macula um V-008141
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
The la e expe on Lilium, P o esso William Thomas S ea n (1911–2001), w i -
ing abou L. japonicum and speci ically abou Thunbe g’s compila ion o Flo a
Japonica, made he ollowing apposi e commen : “His Flo a Japonica whe ein
no less han eigh lilies we e eco ded om Japan, is ull o e oneous de e mi-
na ions; i se ed ne e heless as a basis o la e wo k…” (S ea n 1947: 180).
In he P e ace o his Flo a Japonica, Thunbe g ci ed wo ks connec ed o Ja-
pan and, in pa icula , o h ee ea lie Eu opean bo anis -physicians (Thunbe g
1784: xx i). He men ioned Ch is ian Men zel (1622–1701), who, as pe sonal
physician o F ied ich Wilhelm, Elec o o B andenbu g, would ha e seen he
Honzō kōmoku ansla ed and published in Kyō o by Noda Yajiemon in 1637
and which was kep in he elec o ’s Be lin lib a y. This was he i s Japanese
p in ing o he Chinese Ben Cao Gang Mu [Compendium on Ma e ia Medica],
compiled by he Chinese 16 h-cen u y he balis Li Shizhen (1518–1593), com-
p ising woodcu s o plan s, including species o Lilium, mine als, and animals,
o iginally published pos humously in 1596 in Nanjing, China.
Men zel also ecei ed he 599 colou ed d awings o Japanese plan s which
we e sen back o him by he Ge man And eas Cleye (1634–1698), he oppe -
hoo d o chie me chan on Dejima (Michel 2002: 715). Cleye and his assis an
and ga dene Geo g Meis e (1653–1713) we e pos ed on Dejima wice, i s ly
om 1683 o 1684 and again om 1685 o 1686. Men zel had he illus a ions
bound in o one olume wi h a on ispiece i led Flo a Japanica (Men zel 1695),
which is now in he S aa sbiblio hek in Be lin (shel ma k Lib i pic u a i A 41/42).
Cleye w o e wo sho essays on Japanese lilies ( o mo e de ails, see Lack 2014).
The i s , en i led Obse a io CXCI De Flo ibus Japanensibus, Kanako Ju i e Jama
Ju i (Cleye 1690: 490), includes wo igu es: Cleye 1690: ig. 53, wi h s em bul-
bils in he lea axils and spo ed e lexed lowe s, i led Jama Ju i [moun ain lily],
which ep esen s L. lanci olium Thunb.; and Cleye 1690: ig. 54, i led Kanoko Ju i
[desi able lily], which is eadily iden i iable as L. speciosum Thunb. (Cleye 1690:
490). The second is mo e o less a epea o he i s , en i led Obse a io LXXVI
Flo ibus Japanensibus Vohsno ana & Ghimi (Cleye 1691: 127), accompanied by
Cleye 1691: ig. 20 “Ghimi si e Musme Ju i”. His desc ip ion o i ha ing edible
lea es and bulbs and b ick-colou ed lowe s wi h a e lexed pe igone makes i
iden i iable as L. leich linii Hook. . subsp. maximowiczii (Regel) J.Comp on. Meis-
e also w o e abou lilies in his ea men o Japanese use ul plan s in his wo k
De O ien alisch-Indianische Kuns -und Lus -Gä ne [The O ien al-Indian a and
pleasu e ga dene ], in which he e e ed o wha he called “Sch ogu y Lilium Al-
bum” (i.e. shi o ju i, meaning whi e lily), which was p obably L. longi lo um Thunb.,
and “Cannakogu y” (Cleye ’s kanoko ju i) o L. speciosum (Meis e 1692: 163).
In he same passage o his P e ace in Flo a Japonica, Thunbe g men ioned
Ch is ian Hein ich E ndel (1676–1734), bo anis -physician o F ied ich Augus-
us, Elec o o Saxony (also known as Augus us II he S ong), who published
ex s in 1716 on Japanese medicinal plan s compiled by he Danzig [Gdańsk]
me chan s Jacob B eyne (1637–1697) and his son Johann Philipp B eyne
(1680–1764). As weal hy me chan s, he B eynes’ publica ion was based on
he Japanese collec ions o he VOC and, in pa icula , on hose collec ions and
publica ions o Cleye and Meis e al eady men ioned.
Thunbe g, howe e , singled ou he wo k o his p edecesso on Dejima, he
Ge man physician Engelbe Kaemp e (1651–1716), du ing he la e ’s isi as
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
physician o he VOC on he li le island colony om 1690 o 1692. Kaemp e made
he ho eis o Edo on wo occasions in 1691 and 1692 and la e published his bo-
anical esea ch in Amoeni a um exo ica um (Kaemp e 1712). Thunbe g also a -
emp ed o lea n he Japanese names o he plan s which Kaemp e had included.
Kaemp e included eigh plan s (Kaemp e 1712: 870–872) which he ca e-
go ised as lilies: 1. Sji è, ec ius Sji ói & Osji oi Lilium album, which he called
whi e lily, and judging om his desc ip ion o “Cal hae palus is olio, caule ses-
quicubi ali pingui” meaning lea es like Cal ha palus is L. s em hick, 27 inches
= c. 68 cm all, possibly e e ed o he ela ed lily-like Ca dioc inum co da um
(Thunb.) Makino; 2. Jamma Osji oi, which he desc ip ion s a es is a woodland
plan wi h h ee lea es clasping he s em, could be T illium schonoskii Maxim.;
3. Biakko, ulgo Ju i, wi h scen ed lowe s, which may e e o Polygona um
odo a um (Mill.) D uce a . hunbe gii (C.Mo en & Decne.) H.Ha a; 4. Sazu i
p obably Lilium longi lo um; 5. Ken an, ulgo Oni Ju i, Lilium diaboli, known in
Japan as he de il’s lily Lilium lanci olium; 6. Kasbiako, ulgo Kónnoko Ju i iden-
i iable as L. speciosum; 7. San an, ulgo Fime Ju i, which, om he desc ip ion
o lowe s like a small F i illa ia impe ialis L. bu pu ple-blood ed and spo ed,
migh e e o F i illa ia camscha kensis (L.) Ke Gawl. and 8. Fi Ju i Lilium igne-
um, meaning i e lily, migh e e o L. callosum Siebold & Zucc.
When Thunbe g a i ed in Japan, encoun e ing hi he o unknown Japanese
species o Lilium, hese e e ences, wi h hei equen ly a he inaccu a e de-
pic ions and desc ip ions, we e his p ima y sou ces o consul a ion.
Thunbe g’s i s concep o Lilium o Flo a Japonica
Thunbe g included eigh lilies in his Flo a Japonica (Table 1), which he called:
L. bulbi e um, L. canadense, L. candidum, L. japonicum, L. philadelphicum,
L. pomponicum [sic!], and L. supe bum, as well as wo a ie ies o his new spe-
cies L. japonicum, a s. α and β (Thunbe g 1784: 133–135). He clea ly belie ed
ha Japanese plan s ep esen ing L. bulbi e um and L. candidum L. we e he
same as hei Eu opean coun e pa s, ci ing Linnaeus’s desc ip ions o hose
species e ba im (Linnaeus 1753: 302), including e e ences o Linnaeus’s spe-
cies names in each case. Lilium bulbi e um is con i med as ha ing axilla y bul-
bils along i s s em (Pe uzzi and Ja is 2009), which undoub edly led Thunbe g
o conclude e oneously ha he was obse ing he same species.
Thunbe g also included unde he name L. “pomponicum” he plan wi h ed
Tu k’s cap lowe s ha he belie ed o ep esen he Eu opean L. pomponium L.
(Linnaeus 1753: 303), and ano he as he No h Ame ican L. supe bum L. (Lin-
naeus 1762: 434). As a as Thunbe g’s concep o L. bulbi e um was conce ned,
he ecognised ha he Japanese lily had sca e ed lea es and a campanula e
up igh co olla, bu did no elucida e u he . He ci ed Kaemp e ’s desc ip ion o
he species (Kaemp e 1712: 871) and epea ed Kaemp e ’s Japanese names
o i , “Ken an”, “Oni Ju i”, and “Jamma Ju i” [moun ain lily], bu excluded Kae-
mp e ’s ci a ion o “Lilium diaboli” o de il’s lily (Thunbe g 1784: 134). The e is
a specimen showing bulbils in he lea axils up he s em bu lacking lowe s in
Thunbe g’s he ba ium in Uppsala [UPS-THUNB 8139] wi h an anno a ion w i -
en in Thunbe g’s hand “Lilium lanci olium” and on he e e se “e Japonia Thun-
be g”. This is clea ly iden i iable as L. lanci olium, no L. macula um.
248
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
Thunbe g, howe e , also included he same Japanese names “Ken an”, “Oni
Ju i”, and “Jamma Ju i” o he species, which he called L. canadense on he ollow-
ing page (Thunbe g 1784: 135). Thunbe g again epea ed Linnaeus’s ph ase name
o L. canadense e ba im and s a ed ha his lily had lea es in who ls and e lexed
lowe s, bu also no ed “hinc inde in syl ulis, Nagasaki mon ibus, alibi” [ ound he e
and he e in woods, in he moun ains a ound Nagasaki, and elsewhe e] (Thunbe g
1784: 135). In Thunbe g’s he ba ium in Uppsala, he e is a cul i a ed specimen ha
was anno a ed “Lilium canadense” and on he e e se “cul . in Ho o Uppsaliensis.
Thunbe g” in Thunbe g’s hand [UPS-THUNB 8130]. This specimen has sca e ed
lea es and lowe s wi h na ow, ecu ed epals co e ed wi h many spo s and is
e e able oday o L. leich linii subsp. maximowiczii (Regel) J.Comp on (Comp on
2021). This disc epancy o e names se es o highligh Thunbe g’s ini ial a he
con adic o y de e mina ions o Japanese lilies.
Thunbe g added u he g is o he mill o con usion by including wha he
belie ed o be ano he No h Ame ican lily species, L. philadelphicum, which
had also been desc ibed by Linnaeus (Linnaeus 1762: 435). Thunbe g once
again epea ed Linnaeus’s desc ip i e ph ase name o ha species e ba im,
desc ibing his lily as ha ing who led lea es, up igh lowe s, and pe als wi h
claws (Thunbe g 1784: 135). The e is no specimen in Thunbe g’s he ba ium
wi h his name a ached.
Thunbe g’s second concep o Lilium in Obse a ions on he
Flo a Japonica
Ano he decade passed be o e Thunbe g published he Bo anical Obse a ions
on he Flo a Japonica, which was a commen a y wi h e isions on he species
he had ou lined in his ea lie wo k. The Obse a ions we e ead ou o he Lin-
nean Socie y o London on 1 Oc obe 1793 (Thunbe g 1794: 332–334). Wi h
espec o Lilium, Thunbe g discussed six species (Table 1), one o which, Lil-
ium co di olium Thunb., is now ecognised as belonging in a ela ed genus as
Ca dioc inum co da um, based on his ea lie Heme ocallis co da a Thunb. In
he Obse a ions, ou new species o Lilium we e desc ibed: L. lanci olium,
L. longi lo um, L. macula um and L. speciosum. He also included, as he e-
maining species o he six in he wo k, his second concep o L. bulbi e um,
ci ing as a synonym, he L. philadelphicum, which he had p e iously ci ed in
Flo a Japonica. This was a clea indica ion ha he had changed his mind abou
he iden i y o one o he plan s, which he had o iginally called L. bulbi e um
in Flo a Japonica, e e ing o ha ins ead as a synonym o his new species
L. lanci olium (Thunbe g 1794: 333). Thunbe g’s decision was in luenced by
Kaemp e ’s desc ip ion “gemmis in sinu olio um pisi magni udinis” and “ lo es
…. se iceis dense gu a o” [bulbils in he pocke s o he lea es which a e he
size o peas] and [ lowe s ….. densely clo hed in silky hai s]. Thunbe g ealised
ha his desc ip ion applied equally well o he Eu opean L. bulbi e um as o his
new Japanese species, which also p oduced small bulbils along he s em in he
lea axils. The ige lily, L. lanci olium, known o some ime unde he synonym
L. ig inum Ke Gawl., is now ypi ied on Thunbe g’s shee UPS-THUNB 8139
(Gao and Gao 2017: 174). I is ound h oughou all ou majo islands o Japan,
is widely cul i a ed, and i s diploid o m is only ound on he island o Tsushima,
which lies be ween Kyushu and Sou h Ko ea (Hayashi 2016: 114).
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
In he p o ologue o L. macula um, Thunbe g p o ided he La in diagnosis:
“ oliis spa sis e e icilla is lanceola is, glab is, co ollis campanula is in us
macula is: limbo e lexo.” [lea es spa se, e icilla e, lanceola e, glab ous, co-
ollas campanula e, spo ed wi hin: limb e lexed]. He desc ibed he lea es as
spa se and who led, sessile, lanceola e, glab ous, mul ine ed and inge -like;
he co ollas campanula e wi h he limb e lexed, lesh-pink and inside hey we e
libe ally spo ed wi h many pu ple spo s (Thunbe g 1794: 334). Thunbe g no -
ed ha he s ems we e e e e, s ia ed, hai less, simple, hollow, a oo all and
sub-umbella e a he apex. A e he desc ip ion, he ga e a b ie diagnosis, seg-
ega ing his new lily om he L. canadense Flo . Japon. P. 135, which he had
ea lie e e ed o he Japanese species, i.e. L. leich linii subsp. maximowiczii
(Thunbe g 1784: 135):
“Di e a Lilio canadense: oliis basi la is e co ollis minus e olu is.” [di e s
om Lilium canadense in he b oade basal lea es and he co ollas, which a e
less e olu e].
Also in Thunbe g’s He ba ium in Uppsala is a shee labelled by him “L. mac-
ula um” [UPS-THUNB. 8141] comp ising an in lo escence o wo la ge, up igh
campanula e lowe s wi h spo ing clea ly isible on he epals. This shee has
wo insc ip ions w i en on he ec o in Thunbe g’s hand: “Lilium ma agon” on
he bo om le and “Lilium macula um” on he bo om igh . On he e so, he e
is simply “e Japonia, Thunbe g” also in his hand. This shee is wi hou doub
o iginal ma e ial o he name L. macula um Thunb.
Thunbe g had included in his Flo a Japonica wha he had belie ed was he
No h Ame ican L. philadelphicum (Thunbe g 1784: 135). A decade la e , in his
Obse a ions, he concluded ha his concep o ha species was in ac based
on L. bulbi e um wi h he desc ip ion “ oliis spa sis lanceola is, co ollis campan-
ula is e ec is, caule illoso” [wi h spa sely a anged lanceola e lea es, up igh
campanula e co ollas and illose s ems] (Thunbe g 1794: 333). This desc ip ion
is a sligh ly modi ied e sion o Linnaeus’s p o ologue o L. bulbi e um wi h he
addi ion o “lanceola is” o he lea es and “ illoso” o he s em (Linnaeus 1753:
302). Again, in he Obse a ions, Thunbe g goes on o dis inguish wha he con-
side ed o be he di e ences be ween his second iden i ica ion o L. bulbi e um in
Japan and his ea lie concep o L. philadelphicum (Thunbe g 1794: 334):
“Di e a lilio philadelphico: 1. oliis basi la e io ibus
2. co ollis non e olu is”
[di e s om Lilium philadelphicum: (1). Basal lea es much b oade (2).
Co ollas no e olu e].
In Thunbe g’s He ba ium in Uppsala, he e is ano he shee [UPS-THUNB.
8129] consis ing o wo plan s, bo h wi h lanceola e lea es and each wi h a sin-
gle up igh lowe , nei he o which appea s o ha e spo s on he epals. Each
plan has been ca e ully cu ou om i s o iginal moun and emoun ed on o he
one shee . He clea ly ecognised his as a a ie y o wha he conside ed o be L.
bulbi e um. This shee has wo insc ip ions w i en on he ec o “bulbi e um β l.
Jap” and ‘Lilium bulbi e um d”. On he e so is w i en “e Japonia Thunbe g. Di
a Philad. oliis basi la e io ibus, co ollis e ec is campanula is”. Thunbe g was
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
be ween L. concolo , L. macula um and L. pensyl anicum (Dubouze and Shi-
noda 1999b: 214). Six o hese si es we e sha ed be ween L. macula um and
L. pensyl anicum, while he emaining se en ound in L. macula um could no
be a ibu ed o ei he L. concolo o L. pensyl anicum (Dubouze & Shinoda
Figu e 3. Lec o ype specimen o L. elegans [UPS.THUNB.008129].
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
1999b: 214). Lilium concolo and L. pensyl anicum sha ed six o he 13 si es,
whe eas L. concolo and L. macula um di e ed in all 13 polymo phic si es. In
conclusion, he e was su icien molecula di e gence be ween L. macula um
and he closely ela ed L. pensyl anicum o main ain L. macula um as a sepa-
a e species (Dubouze and Shinoda 1999b: 217). They did, howe e , ask he
highly pe inen ques ion – “jus how many nucleo ide di e ences a e equi ed
o con i m he dis inc ions be ween sec ion, species and subspecies in Lilium?”
(Dubouze and Shinoda 1999b: 217). They implied ha he numbe o base di -
e ences is di ec ly p opo ional o he le el o gene ic di e gence and ha ,
Figu e 4. Ignaz Klaube ’s eng a ings in Memoi s o he Academy o S . Pe e sbu g, ol. 3 (1811), o Johan Olin’s illus a-
ions o L. lanci olium and L. elegans based on Thunbe g’s collec ions in UPS.
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
logically, in aspeci ic a ia ion could be ca ego ised by such axa ha ing ewe
han se en polymo phisms (Dubouze and Shinoda 1999b: 217). They s a ed
ha i was qui e possible ha popula ions o L. macula um we e he emnan s
on Honshu o a sou he ly sp ead om Hokkaido o L. pensyl anicum, and ha
hose con inued o specia e a e he las glacial pe iod, i.e. c. 10,000 ybp. They
posi ed ha hese popula ions de eloped mechanisms such as immedia e epi-
geal ge mina ion ha allowed hem o compe e and su i e in he milde , mo e
sou he ly clima e o Honshu (Dubouze and Shinoda 1999b: 217).
Molecula phylogenies including hese h ee species based on bo h nuclea
and plas id DNA, ha e shown ha L. macula um belongs ei he on a clade wi h
L. bulbi e um and L. pensyl anicum (sampled as L. dau icum) sis e o a clade
wi h L. concolo (Nishikawa e al. 2001: 43) o on a clade wi h L. lanci olium and
L. pensyl anicum sis e o a clade wi h L. callosum, L. concolo and L. leich linii
(Nishikawa e al. 1999: 248; Dubouze and Shinoda 1999a: 957; Dubouze and
Shinoda 1999b: 217; Gi nish e al. 2020: 39). Addi ional s udies ha e shown
ha L. pensyl anicum (as L. dau icum) si s on a b anch wi h L. leich linii sis e
o L. lanci olium and L. macula um (Du e al. 2014: 254) o ha L. macula um
and L. macula um a . bukosanense o med a b anch sis e o L. da idii wi h
Figu e 5. Lilium macula um g owing nea he eas coas along Thunbe g’s ou e pas
Omaezaki, Shizuoka P e . Pho o. Skyca . 2 June 2021.
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
weak boo s ap suppo (68% maximum pa simony boo s ap and 62% max-
imum likelihood boo s ap) while L. pensyl anicum (as L. dau icum) o med a
e y s ongly suppo ed b anch wi h L. lanci olium a . la i lo um Makino (98%
MP and 98% ML), sis e o L. leich linii Hook. . (Wa anabe e al. 2021: 192).
In none o hese phylogenies is i e iden ha L. macula um and L. concolo
occu on he same b anch o he ele an clade, no is he e any e idence om
mo phology ha L. macula um exhibi s any cha ac e s linking he wo species.
All he axa men ioned he e belong o an amended Lilium sec . Sinoma a-
gon H.F.Combe (Combe 1949: 101), when i is combined wi h he mono ypic
L. sec . Dau oli ion H.F.Combe , whose sec ional name was based solely on
L. pensyl anicum (as L. dau icum) (Combe 1949: 102). Dubouze and Shinoda
(1999b: 217) concluded ha L. macula um and L. pensyl anicum a e closely
ela ed bu disc e e species, ha L. concolo is sis e o hem bo h, and ha
L. macula um is no o hyb id o igin.
Lilium ×elegans
Thunbe g’s Lilium elegans has been sugges ed o be a hyb id be ween L. pen-
syl anicum and L. macula um (Shimizu 1960: 105; 1971: 277; 1987: 75; Fu uya
2009: 85). The e y la ge ange o highly o namen al cul i a ed ma e ial ound
in Japan b ed o e many cen u ies has been he basis o his sugges ion. Ka y-
ological esea ch ci ed by Shimizu (1960: 115) indica ed ha he ch omosomes
o L. macula um di e ed om hose o L. pensyl anicum (as L. dau icum). The
la e we e shown o ha e seconda y cons ic ions in he se en h pai o ch o-
mosomes, whe eas hose o L. macula um lacked any seconda y cons ic ions
(Kumazawa and Kimu a 1946). A mo e ecen ka yological s udy on 32 species
o Lilium in China ound ha he a ia ion in ch omosome mo phology ac oss
he genus in China as a whole is ela i ely low (Gao e al. 2011: 749). Lilium
macula um, howe e , was no sampled as ha s udy was es ic ed solely o
Chinese axa. Howe e , he sample o L. pensyl anicum (as L. dau icum) in-
cluded by Gao e al. (2011) did no e eal he p esence o seconda y cons ic-
ions in he se en h pai o ch omosomes (Gao e al. 2011: 758). Shimizu, when
discussing he mo phology o he pu a i e pa en species o L. ×elegans, i.e.
L. pensyl anicum and L. macula um, wen on o dis inguish L. macula um mo -
phologically by i s p oduc ion o unde g ound s olons which can each mo e
han 10 cm in leng h be o e eme ging om he soil (Shimizu 1960: 111). This
ea u e, howe e , was used by Hayashi o e e o L. pensyl anicum and o dis-
inguish i om L. macula um in his ea men o Lilium in he Flo a o Japan
(Hayashi 2016: 112). I is likely, howe e , ha bo h species exhibi unde g ound
s olon o ma ion when he condi ions o s olon p oduc ion a e a ou able, i.e.
when plan s a e g owing in e ile, ee-d aining, loamy soil.
F om he in o ma ion p o ided by Thunbe g (1811: 203) and examina ion o
his he ba ium ma e ial o L. elegans, UPS-THUNB. 8129 (Fig. 3), i is e iden
ha he was desc ibing a plan ha a ied om L. macula um. The e is no e -
idence as o exac ly whe e Thunbe g may ha e acqui ed his ma e ial, bu as
i had been long-cul i a ed in Japan, he assump ion mus be ha i was o
ga den o igin. The ques ion o he a ibu ion o he co ec name o any hyb id
be ween L. pensyl anicum and L. macula um would he e o e be conside ed
he e. I is well ecognised in Japanese li e a u e ha plan s ha e been b ed
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
om hese species o hund eds o yea s. In wes e n li e a u e, i was egula ly
hough ha L. hunbe gianum was he co ec name o be used o his hyb id;
howe e , L. elegans was desc ibed in 1811 and L. hunbe gianum in 1829 and
included he ea lie name in synonymy. The name L. hunbe gianum is he e o e
conside ed o be an illegi ima e name (A . 52.1 o he ICN) au oma ically ypi-
ied (A . 7.5 o he ICN) by he ype o he name L. elegans.
The e idence o hyb idisa ion be ween L. macula um and L. pensyl anicum
is ex emely well p esen ed based on e idence om sequence da a om i e
plas id genes and wo nuclea genes (Wa anabe e al. 2024: 458, 459). Howe -
e , hey did no ci e a alid hyb id name, A . H3.1 s a es ha hyb ids be ween
wo o mo e axa may ecei e a name, and H10.1 s a es ha names o no ho-
axa (hyb ids) mus con o m o he p o isions o he Code. A 10.1 No e 1
s a es ha axa p e iously published as species may be indica ed as hyb id
axa wi hou a change o ank. The i s a ailable name o a hyb id be ween he
widesp ead L. pensyl anicum and he mo e es ic ed Japanese L. macula um
is L. ×elegans Thunb. This name can be applied o all ho icul u al cul i a s o
hyb id o igin wi h his pa en age.
Na u al a ia ion wi hin L. macula um
Wa anabe e al. (2024) ecognise wo species and ou in aspeci ic axa
wi hin hei concep o L. macula um as well as he hyb id be ween L. mac-
ula um and L. pensyl anicum. They ega d L. macula um a . macula um as
occu ing along he eas Tōhoku coas on he no h-eas e n pa o Honshu
and on he Sado-Tobishima islands o he wes coas o Honshu (Wa a-
nabe e al. 2024: 463, 464). They added ha L. macula um occu s in coas -
al a eas o Niiga a P e ec u e o he Chūbu dis ic acing he Japan Sea
(Wa anabe e al. 2024: 465). These places lie along he no he n and wes -
e n pa s o Honshu and would ha e been a om he a eas whe e Thun-
be g was pe mi ed o a el in o de o collec his lilies du ing he ho eis o
Edo in he summe o 1776. Indeed, his ou e lay along he Paci ic coas o
sou he n Honshu (Kijewski 2015: 162). In e ec , Thunbe g simply could no
ha e seen L. macula um om hose no he n and wes e n egions o Japan.
Wa anabe e al. claimed ha L. macula um, which hey say is some imes
called in Japanese, “Iwa-yu i” [ ock lily], occu s on he coas o Wes Tōhoku
in no he n Honshu and includes wha hey conside ed o be he ype o L.
macula um (Wa anabe e al. 2024: 465). Mo eo e , hey also claimed ha L.
macula um had o igina ed gene ically by in og ession as pa o hei new
species L. paci icum S.T.Wa an., Fuse & M.N.Tamu a, whose holo ype came
om he Izu peninsula in Shizuoka P e ec u e along he Paci ic coas (Wa a-
nabe e al. 2024: 465). Al hough he e is no e idence ha Thunbe g isi ed
he peninsula, i is close o his ou e o Edo along he Paci ic eas coas .
Wa anabe e al. (2024: 465) dis inguished wha hey unde s ood o be L. mac-
ula um om hei L. paci icum by he p esence o papillose hai s on he mid ibs
o he adaxial lea su aces o L. macula um ( s. smoo h on L. paci icum). They
desc ibed L. paci icum om he eas coas o Shizuoka P e ec u e, and as oc-
cu ing in coas al pa s o Kan o and eas e n Chūbu dis ic s o Honshu. These
a e ce ainly he egions ha Thunbe g had passed h ough on his ho eis
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James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
jou ney bo h o and om Edo (Kijewski 2015). Thunbe g’s o iginal ma e ial o L.
macula um in UPS [UPS-THUNB 8141] p esen s no e idence o papillose mid-
ibs along he mid eins on he adaxial lea su aces. The ma e ial desc ibed by
Wa anabe e al. as L. paci icum (Wa anabe e al. 2024: 466), lacking papillae
and occu ing om one o he egions a e sed by Thunbe g and ma ching
Thunbe g’s own o iginal ma e ial, he e o e ep esen s L. macula um (Fig. 5).
Conside ing commen s conce ning he pu a i e ecen specia ion o L. mac-
ula um om a elic popula ion o L. pensyl anicum ex ending sou h in o no h-
e n Honshu (Dubouze and Shinoda 1999b: 217) and he commen s by Wa a-
nabe e al. (2024: 465) conce ning he high deg ee o gene low by in og ession
wi hin L. macula um, i seems sensible o ecognise he axon hey unde s ood
as L. macula um om coas al egions nea he Japan Sea on he wes side o
Honshu as a dis inc axon. In addi ion, he le els o a ia ion wi hin L. macula-
um ac oss he ange o he species, as shown by Wa anabe e al. (2024: 465),
me i ecogni ion o he ma e ial desc ibed by hem (as L. macula um) a he
ank o a ie y. Wa anabe e al. (2024: 466) ha e al eady desc ibed he ma e ial
om Sado Island and Niiga a P e ec u e on he wes coas o Japan as L. mac-
ula um a . sadoense S.T.Wa an., Fuse & M.N.Tamu a; consequen ly, his name
is aken up he e o he a ian wi h lea es possessing papillae on adaxial (and
abaxial) lea su aces.
Iden i ica ion key o dis inguish L. concolo , L. ×elegans, L. macula um
and L. pensyl anicum
1 Flowe s s ella e in ou line, pe igone segmen s usually ed, some imes
spo ed, ( a ely yellow in L. concolo a . co idion (Siebold and de V iese)
Bake ) 2.5–6 × 0.5–1.5 cm .......................................................... L. concolo
– Flowe s bowl-shaped in ou line, pe igone segmen s 7–10 × 1.5–2.5 cm,
plain o spo ed ..............................................................................................2
2 Plan s obus , e ec , s em base ei he papillose o smoo h, in lo escence
umbella e, lowe s (3–)5–20, o ange, ed, pink o yellow .......... L. ×elegans
– Plan s e ec o decumben , in lo escence sub-umbella e, lowe s 1–5(–
10), o ange o ed, a ely yellow ....................................................................3
3 S ems glab ous on lowe po ion, ibbed, loccose wi h long hai s nea
apex, densely so in lea axils, lowe buds and a he base o epal idges .
.............................................................................................L. pensyl anicum
– S ems papillose scab id a base, no ibbed, epal idges glab ous ............
...................................................................................................L. macula um
Iden i ica ion key o dis inguish he a ie ies o L. macula um
1 S ems hin, laxly decumben , lea es linea , 0.4–1.2 cm wide, papillose on
mid ib, lowe s 1 (–2) .................................................................................... 2
– S ems s ou , e ec , lea es lanceola e 1–2 cm wide ....................................3
2 S igma o ange ............................................L. macula um a . bukosanense
– S igma eddish-b own ..................................... L. macula um a . mon icola
3 Adaxial lea mid ibs papillose .........................L. macula um a . sadoense
– Adaxial lea mid ibs glab ous ....................... L. macula um a . macula um
262
Phy oKeys 263: 241–270 (2025), DOI: 10.3897/phy okeys.263.163611
James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
Synonymic conspec us
Lilium macula um Thunb., T ans. Linn. Soc. London 2: 334 (1794). Holo ype:
Japan, “e Japonia Thunbe g, Lilium macula um”, C.P. Thunbe g s.n. (holo.
UPS -THUNB. 8141!) [V-008141].
= Lilium elegans a . spon aneum Makino, J. Jap. Bo . 8: 40 (1932) ≡ L. mac-
ula um . spon aneum (Makino) S.T.Wa an., Fuse & M.N.Tamu a, Taxon
73(2): 465 (2024). Lec o ype designa ed by Wa anabe e al. 2024: Japan,
Iwa e P e ., Omoe Peninsula, 11–13 Sep embe 1929, T. Makino s.n. (lec-
o. MAK 139176).
= Lilium paci icum S.T.Wa an., Fuse & M.N.Tamu a, Taxon 73(2): 466 (2024).
Holo ype: Japan, Shizuoka P e . Shimoda-shi, 21 June 1979 ( l.), S. Ki amu a
193 (KYO holo!) [KYO00029149] iso ypes: [KYO00029150; KYO00029151].
Lilium macula um a . bukosanense (Honda) H.Ha a, J. Jap. Bo . 38(8): 249
(1963). ≡ Lilium bukosanense Honda in Nakai, Iconog . Pl. Asiae O ien . [Toa
Shokubu su Zuse su] Tokyo 4(3): 404 . 128 (1942). Holo ype: Japan, Hon-
shu, Musasi [Chichibu, Sai ama P e ec u e], “Musashi no Kune, in mon e
Bukōsan, Shimizu Daisuke s.n. 18 July 1941” (TI holo.!) [TI00083088] Japa-
nese name: Miyama Sukashi Yu i
= Lilium macula um . la e i ium Nakai, J. Jap. Bo . 18: 164 (1942). Holo ype:
Japan, Tokyo, Aogashima Island, 7 July 1932, M. Takahashi s.n. (holo. TI)
[TI00083089].
Lilium macula um a . mon icola H.Ha a, J. Jap. Bo . 38(8): 249 (1963)
≡ L. macula um . mon icola (H.Ha a) S.T.Wa an., Fuse & M.N.Tamu a, Taxon
73(2): 465 (2024). Holo ype: Cul . Tokyo Bo anic Ga den H.Ha a s.n, 14 June
1962. Japan, Honshu, Yamaga a P e ., A agoyama (TI) [TI00083090]
Lilium macula um a . sadoense S.T.Wa an., Fuse & M.N.Tamu a, Taxon 73(2):
466 (2024). Holo ype: Japan, Niiga a P e ., Sado Island, Aikawaou a, ocky
seaside al . 10 m., 7 June 1991, K. Hayashi 80 (KYO holo!) [KYO00029152]
iso ypes; [KYO00029153, KYO00029155].
Lilium ×elegans Thunb., Mém. Acad. Imp. Sci. S . Pé e sbou g His . Acad. 3:
203 (1811) ≡ L. macula um a . elegans (Thunb.) Koidz., Bo . Mag. (Tokyo)
39: 307 (1925) ≡ L. hunbe gianum Schul . & Schul . ., Sys . Veg. ed. 15bis
7(1): 415 (1829) [L. bulbi e um Thunb. “di e a L. philadelphico. In Japonia”
was ci ed] nom. illeg. (A . 52.1 o he ICN) ≡ L. bulbi e um subsp. hunbe -
gianum (Schul . & Schul . .) Bake nom. illeg., Ga d. Ch on. 1871(2): 1034
(1871) ≡ L. bulbi e um a . hunbe gianum (Schul . & Schul . .) Bake nom. il-
leg., J. Roy. Ho . Soc. [n.s.] 4(1): 43 (1873). ≡ L. dau icum subsp. hunbe gia-
num (Schul . & Schul . .) E.H.Wilson nom. illeg., Lilies Eas Asia: 53 (1925).
Lec o ype designa ed by Wa anabe e al. 2024: 469: “e Japonia, Thunbe g,
di . a Philad. oliis basi la io ibus, co ollis e ec is campanula is”, “bulbi e um
β Fl. Jap.”, C.P.Thunbe g s.n. (lec o.! UPS -THUNB. 8129) [V-008129].
= Lilium ulgens C.Mo en in [Eds. Pie e Co neille Van Géel & Pie e Joseph
Augus e D apiez] Encyclog aphie du Règne Végé al p . 6 sui e du gen e Lis
– Lilium ulgens . 10 (1835) ≡ L. bulbi e um a . ulgens (C.Mo en) Bake ,
263
Phy oKeys 263: 241–270 (2025), DOI: 10.3897/phy okeys.263.163611
James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
J. Roy. Ho . Soc. [n.s.] 4(1): 44 (1873) ≡ L. bulbi e um . ulgens (C.Mo en)
A.Wallace, No es Lilies: 71 (1873) ≡ L. elegans a . ulgens (C.Mo en) Bake ,
J. Linn. Soc. Bo . 14: 240 (1875). Lec o ype (designa ed he e): [Icon] Ency-
clog aphie du Règne Végé al p . 6 sui e du gen e Lis – Lilium . 10 (1835)
= Lilium enus um F.Neumann, Allgemeine Deu sche Ga en-Zei ung 18(20):
157 (16 May 1840). (see No e unde L. enus um Kun h below).
= Lilium enus um Kun h, Enum. Pl. 4: 265 (1843) ≡ L. bulbi e um . enus um
(Kun h) A.Wallace, No es Lilies: 71 (1873) ≡ L. hunbe gianum a . enus um
(Kun h) Maxim. ex F anch & Sa ., Enum. Pl. Jap. 2: 69 (1879) ≡ L. elegans
a . enus um (Kun h) Elwes, Monog . Lilium 5: . 19 (1878) ≡ L. dau icum [as
da u icum] a . enus um (Kun h) E.H.Wilson, Lilies E. Asia: 52 (1925).
No e 1: Lilium enus um F.Neumann, Allgemeine Deu sche Ga en-Zei ung
18(20): 157 (16 May 1840) p eda es L. enus um Kun h, Enum. Pl. 4: 265
(1843). Neumann’s desc ip ion s a es: “Eine seh schöne Lilie, die den Bau
de gemeinen Feue lilie (L. bulbi e um) ha , sich abe on ih un e schei-
de : 1. dass die Blumen eine blassgelbe Fa be haben, ohne alle Punk e
und Fleke; 2. dass die Blumenblä e mi ih en Rände n an einande geleg
bleiben, 3. dass die Blumen sich seh lange hal en, weshalb sie om Juli
bis im Sep embe o blüh und endlich 4. dass sie keine Bulbillen in den
Bla winkeln äg , weshalb ih e Ve meh ung nu langsam o sich geh ”
ansla ed as “A e y beau i ul lily, ha ing he s uc u e o he gemmed
i e lily (L. bulbi e um), bu di e ing om i : 1. ha he lowe s a e o a
pale-yellow colou , wi hou any spo s o speckling; 2. ha he pe als si
wi h hei edges oge he , 3. ha he lowe s las o a e y long ime,
which is why i con inues o bloom om July o Sep embe and inally 4.
ha i has no bulbils in he axils o he lea es, which is why i will ep o-
duce only slowly”. Lilium enus um Kun h, Enum. Pl. 4: 265 (1843) is de-
sc ibed as ha ing lowe s: “in undibula i-campanula is, au an iaco-mini-
a is colo ibus” i.e. wi h o ange-sca le lowe s. This is clea ly hen based
on a di e en ype and is o be conside ed an illegi ima e name A . 52.1.
The ollowing combina ions howe e , a e alid (see synonymy abo e):
L. bulbi e um . enus um (Kun h) A.Wallace; L. elegans a . enus um
(Kun h) Elwes; L. hunbe gianum a . enus um (Kun h) Maxim. ex F anch
& Sa . and L. dau icum [as da u icum] a . enus um (Kun h) E.H.Wilson.
= Lilium sanguineum Lindl., Edwa ds’s Bo . Reg. 32 [n.s. 9] . 50 (1846). ≡ L.
hunbe gianum . sanguineum (Lindl.) Bake & Dye , Ga d. Ch on. 1872: 1356
(1872) ≡ L. bulbi e um . sanguineum (Lindl.) A.Wallace, No es Lilies: 71
(1873) ≡ L. bulbi e um a . sanguineum (Lindl.) Bake , J. Roy. Ho . Soc., [n.s.]
4(1): 43 (1873) ≡ L. elegans a . sanguineum (Lindl.) Bake , J. Linn. Soc. Bo .
14: 239 (1875) ≡ L. dau icum [as da u icum] . sanguineum (Lindl.) E.H.Wil-
son, Lilies E. Asia: 55 (1925). Lec o ype designa ed by Wa anabe e al. 2024:
468: [icon] Edwa ds’s Bo . Reg. 32 [n.s. 9] . 50 (1846)
= Lilium ulgens a . macula um Spae, Mém. Cou onnés Mém. Sa an s E ang-
e s Acad. Roy. Sci. B uxelles 19(5): 20 (1847). Type: Siebold ex Japan s.n.
1825–1830 l. Ghen Bo anic Ga den 1833 (no seen).
= Lilium ulgens a . s aminosum Lem., Ill. Ho . 11: . 422 (1864). Lec o ype
designa ed he e: [Icon] “In ho o Ve scha el , P. S ooban pinxi ” Ill. Ho .
11: . 422 (1864).
264
Phy oKeys 263: 241–270 (2025), DOI: 10.3897/phy okeys.263.163611
James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
= Lilium o mosum Lem., Ill. Ho . 12: . 459 (1865). Lec o ype designa ed he e:
[icon] “P. S ooban ad na . pinxi . In ho o Ve scha el . Japon (plein ai )”, Ill.
Ho . 12: . 459 (1865).
= Lilium haema och oum Lem., Ill. Ho . 14: 503 (1867) ≡ L. bulbi e um . hae-
ma och oum (Lem.) A.Wallace, No es Lilies: 71 (1873). Lec o ype designa -
ed by Wa anabe e al. 2024: 469: [Icon] “In ho o Ve scha el , P. S ooban
pinxi , Japon ai lib e” Ill. Ho . 14: 503 (1867)
= Lilium hunbe gianum a . pa dinum T.Moo e, Flo is & Pomologis se . 3(1):
121 (1868) ≡ L. ×wilsonii Leich lin ex R.Hogg, T.Moo e & W.Paul, Flo is &
Pomologis se . 3(1): 192 (1868) ≡ L. bulbi e um . wilsonii (Leich lin ex
R.Hogg, T.Moo e & W.Paul) A.Wallace, No es Lilies: 70 (1873) ≡ L. bulbi-
e um a . wilsonii (Leich lin ex R.Hogg, T.Moo e & W.Paul) Bake , J. Roy.
Ho . Soc. [n.s.] 4(1): 43 (1873) ≡ L. elegans a . pa dinum (T.Moo e) Bak-
e , J. Linn. Soc. Bo . 14: 239 (1875) ≡ L. dau icum [as da u icum] . pa di-
num (T.Moo e) E.H.Wilson, Lilies E. Asia: 52 (1925). Lec o ype designa ed
by Wa anabe e al. 2024: 470: [icon] Flo is and Pomologis 1868: 121 .
“Lilium hunbe gianum pa dinum” (1868).
No e 2: Lilium ×wilsonii Leich lin ex R.Hogg, T.Moo e & W.Paul.
As s a ed in he p o ologue o L. hunbe gianum [un anked] pa dinum, his
is o hyb id o igin published by Thomas Moo e (1821–1887) in he June issue
o Flo is and Pomologis o he yea 1868. The new combina ion Lilium ×wil-
sonii published in he same olume bu in Augus ha same yea , is a enaming
o he un anked in aspeci ic name published ea lie in June. The a he b oad
lea es wi h i e longi udinal eins and he da k papillose ma kings wi hin he
basal pa s o each epal, combined wi h he deep yellow zone along he sinus,
sugges he possibili y ha L. au a um Lindl. may ha e c ossed wi h he up igh
bowl-shaped eddish-o ange lowe s o L. macula um.
= Lilium al e nans Siebold ex Duch., J. Soc. Na l. Ho . F ance se . 2 ol. 4 (4):
215, 473 (1870).
= Lilium hunbe gianum a . bicolo T.Moo e, Flo al Mag. 9: . 504 (1870) ≡ L. bul-
bi e um . bicolo (T.Moo e) A.Wallace, No es Lilies: 70 (1873) ≡ L. bulbi e um
a . bicolo (T.Moo e) Bake , J. Roy. Ho . Soc., [n.s.] 4(1): 43 (1873) ≡ L. ele-
gans a . bicolo (T.Moo e) Bake , J. Linn. Soc. Bo . 14: 239 (1875). Lec o ype
designa ed by Wa anabe e al. 2024: 469: [icon] Flo al Mag. 9: . 504 (1870).
= Lilium hunbe gianum a . a osanguineum Bake & Dye , Ga d. Ch on. 1872:
1356 (1872) ≡ L. bulbi e um . a osanguineum (Bake & Dye ) A.Wallace, No es
Lilies: 71 (1873) ≡ L. bulbi e um a . a osanguineum (Bake & Dye ) Bake , J.
Roy. Ho . Soc., [n.s.] 4(1): 43 (1873) ≡ L. elegans a . a osanguineum (Bake
& Dye ) Bake , J. Linn. Soc. Bo . 14: 240 (1875) ≡ L. dau icum [as da u icum] .
a osanguineum (Bake & Dye ) E.H.Wilson, Lilies E. Asia: 55 (1925).
= Lilium hunbe gianum . b e i olium Bake & Dye , Ga d. Ch on. 1872: 1356
(1872) ≡ L. bulbi e um a . b e i olium (Bake & Dye ) Bake , J. Roy. Ho .
Soc., [n.s.] 4(1): 43 (1873) ≡ L. elegans a . b e i olium (Bake & Dye ) Bake ,
J. Linn. Soc. Bo . 14: 239 (1875)
= Lilium hunbe gianum . alu aceum Bake & Dye , Ga d. Ch on. 1872: 1356
(1872) ≡ L. bulbi e um . alu aceum (Bake & Dye ) A.Wallace, No es Lilies:
70 (1873) ≡ L. bulbi e um a . alu aceum (Bake & Dye ) Bake , J. Roy. Ho .
265
Phy oKeys 263: 241–270 (2025), DOI: 10.3897/phy okeys.263.163611
James A. Comp on: A eapp aisal o Thunbe g’s spo ed lily Lilium macula um Thunb.
Soc., [n.s.] 4(1): 43 (1873) ≡ L. elegans a . alu aceum (Bake & Dye ) Bake , J.
Linn. Soc. Bo . 14: 239 (1875) ≡ L. dau icum [as da u icum] . nig o-macula-
um E.H.Wilson, Lilies E. Asia: 52 (1925) ≡ L. macula um . alu aceum (Bake
& Dye ) Nakai, J. Jap. Bo . 18: 166 (1942). Lec o ype designa ed by Wa a-
nabe e al. 2024: 468: [icon] “L. hunbe gianum au eum nig o-macula um Van
Hou e” Fl. Se es 16: . 1627 (1865).
= Lilium hunbe gianum . a meniacum Bake & Dye , Ga d. Ch on. 1872: 1356
(1872) ≡ L. bulbi e um . a meniacum (Bake & Dye ) A.Wallace, No es Lilies:
71 (1873) ≡ L. bulbi e um a . a meniacum (Bake & Dye ) Bake , J. Roy. Ho .
Soc., [n.s.] 4(1): 43 (1873) ≡ L. elegans a . a meniacum (Bake & Dye ) Bake ,
J. Linn. Soc. Bo . 14: 239 (1875).
= Lilium hunbe gianum . b e i olium Bake & Dye , Ga d. Ch on. 1872: 1356
(1872) ≡ L. bulbi e um . b e i olium (Bake & Dye ) A.Wallace, No es Lilies:
70 (1873) ≡ L. bulbi e um a . b e i olium (Bake & Dye ) Bake , J. Roy. Ho .
Soc., [n.s.] 4(1): 43 (1873) ≡ L. dau icum [as da u icum] . b e i olium (Bake
& Dye ) E.H.Wilson, Lilies E. Asia: 55 (1925).
= Lilium dau icum [as da u icum] a . c oceum Regel, Ga en l. 21: 295 (1872)
= Lilium hunbe gianum a . au an iacum Regel, Ga en l. 21: 295 (1872)
= Lilium hunbe gianum a . s amineum Regel, Ga en l. 21: 295 (1872) ≡ L. ele-
gans a . s amineum (Regel) Voss, Vilm. Blumengä . ed. 3 ol. 1: 1099 (1896).
= Lilium hunbe gianum a . leich linii Regel, Ga en l. 21: 295 (1872)
= Lilium hunbe gianum . ci inum Bake & Dye , Ga d. Ch on. 1872: 1356 (1872)
≡ L. bulbi e um . ci inum (Bake & Dye ) A.Wallace, No es Lilies: 71 (1873)
≡ L. bulbi e um a . ci inum (Bake & Dye ) Bake , J. Roy. Ho . Soc., [n.s.]
4(1): 43 (1873) ≡ L. elegans a . ci inum (Bake & Dye ) Bake , J. Linn. Soc.
Bo . 14: 239 (1875) ≡ L. dau icum [as da u icum] . ci inum (Bake & Dye )
E.H.Wilson, Lilies E. Asia: 55 (1925).
= Lilium elegans a . incompa abile Elwes, Monog . Lilium 5: . 20 (1878). Lec o-
ype designa ed by Wa anabe e al. 2024: 469: [icon] Elwes, Monog . Lilium
5: . 20 (1878).
= Lilium ba emanniae A.Wallace, The Ga den 15: 396 (1879) ≡ L. elegans a .
ba emanniae (A.Wallace) Bake , J. Roy. Ho . Soc. [London] [n.s.] 26: 339
(1901–1902) ≡ L. dau icum [as da u icum] . ba emanniae (A.Wallace)
E.H.Wilson, Lilies E. Asia: 52 (1925). Lec o ype: designa ed by Wa anabe e
al. 2024: 469 [Icon] The Ga den 15: 396, M s Ba eman’s Lily . clxxx (1879).
= Lilium dau icum [as da u icum] . plu i lo um Voss, Vilm. Blumengä . ed. 3
ol. 1: 1097 (1896).
= Lilium wallacei R.W.Wallace, The Ga den 51: 80, . 1103 (1897) ≡ L. elegans a .
wallacei (R.W.Wallace) Waugh, Cycl. Am. Ho . 2(c3): 919 (1900) ≡ L. dau i-
cum [as da u icum] a . wallacei (R.W.Wallace) E.H.Wilson, Lilies E. Asia: 52
(1925). Lec o ype designa ed by Wa anabe e al. 2024: 469: [icon] The Ga -
den 51: 80, . 1103 (1897).
= Lilium dau icum [as da u icum] . au eum E.H.Wilson, Lilies E. Asia: 55 (1925).
= Lilium dau icum [as da u icum] . ho smannii E.H.Wilson, Lilies E. Asia:
55 (1925).
- Lilium au an iacum Pax on, Pax on Bo . Mag. 6: 127 (1839) nom. illegi .
(L. au an iacum Wes on Bo . Uni . 3: 453. 1772 = L. bulbi e um L.) A . 52.1.
- Lilium a osanguineum H.Vilm., Fl. Pleine Te e ed. 3: 629 (1870) nom.
nud. in syn.
