281
Resol ing he axonomy o Leea i olia a (Leeaceae, Vi ales):
Reins a emen and ypi ica ion
Am u ha A halappil1,2 , Ayillia h Ku iye i P adeep1, Jun Wen2, Nikhil K ishna1,3
1 Depa men o Bo any, Uni e si y o Calicu , Ke ala 673 635, India
2 Depa men o Bo any, Na ional Museum o Na u al His o y, Smi hsonian Ins i u ion, Washing on, DC, 20013-7012, USA
3 Regional Ayu eda Resea ch Cen e, Dimapu , Nagaland 797112, India
Co esponding au ho s: Ayillia h Ku iye i P adeep ([email p o ec ed]); Jun Wen ([email p o ec ed])
Copy igh : This is an open access a icle
dis ibu ed unde he e ms o he CC0 Public
Domain Dedica ion.
Resea ch A icle
Abs ac
Leea i olia a M.A.Lawson has long been conside ed conspeci ic wi h L. compac i lo a
Ku z. De ailed examina ion o li e ma e ials and he ba ium specimens sugges s ha
bo h species a e dis inc , di e ing in habi , lea a chi ec u e, and in lo escence and b ac
mo phology. This con ibu ion p esen s mo phological e idence o he eins a emen
o L. i olia a, including i s axonomic desc ip ion, illus a ion, dis ibu ion maps, no es
on habi a , and conse a ion s a us assessmen using he IUCN c i e ia. In addi ion,
A neo ype was selec ed o L. compac i lo a.
Key wo ds: IUCN, Leea, neo ype, Old Wo ld opics, Vi aceae
In oduc ion
The economically impo an g ape o de Vi ales has wo amilies, Leeaceae
and Vi aceae (Wen 2007a, 2007b), which ha e some imes been ecognized
as sub amilies Leeoideae Bu meis e and Vi oideae Ea on o Vi aceae (APG III
2009; APG IV 2016). The o de is widely dis ibu ed in he opical and empe a e
zones, wi h Leeaceae being mono ypic, consis ing o he genus Leea D.Royen ex
L. Many axonomis s (e.g. Planchon 1887; Gagnepain 1950; Suessengu h 1953;
Ridsdale 1974, 1976; Nai hani 2000; La i 2001; Ren e al. 2003; Chen and Wen
2007; Wen 2007a) ha e long ea ed Leea in i s own amily Leeaceae, due o i s
usually sh ubby o ee habi , absence o end ils, p esence o pe iola s ipules,
de elopmen o a complex lo al disc a ached o a polype alous co olla, he
absence o nec a p oduc ion, he p esence o seconda y sep a in he o a y, he
locules o he o a y each ha ing only one o ule, as well as he dis inc wood and
palynological cha ac e s. The simila i y o Leea and membe s o Vi aceae has
been no ed in e ms o seed and emb yo cha ac e s as well as he occu ence
o pea l glands on he ege a i e o gans (Ridsdale 1974; Wen 2007a, 2007b).
Leea consis s o c. 45 species (POWO 2025) and is dis ibu ed h ough-
ou he Old Wo ld opics, anging om A ica and Madagasca o Fiji (Rids-
dale 1976). The Malesian A chipelago has he highes numbe o species
and endemism, wi h 25 species in his egion (Ridsdale 1974, 1976; Lok e al.
Academic edi o : Anna T ias-Blasi
Recei ed:
30 Sep embe 2025
Accep ed:
10 No embe 2025
Published:
21 No embe 2025
Ci a ion: A halappil A, P adeep AK,
Wen J, K ishna N (2025) Resol ing
he axonomy o Leea i olia a
(Leeaceae, Vi ales): Reins a emen
and ypi ica ion. Phy oKeys 266:
281–295. h ps://doi.o g/10.3897/
phy okeys.266.173605
Phy oKeys 266: 281–295 (2025)
DOI: 10.3897/phy okeys.266.173605
282
Phy oKeys 266: 281–295 (2025), DOI: 10.3897/phy okeys.266.173605
Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
2011). Wi hin he genus, he ci cumsc ip ion o many o he species emains
con o e sial (Molina e al. 2013), and he only signi ican axonomic wo k
on he genus is he e ision o Leeaceae by Ridsdale (1974, 1976). Howe e ,
Ridsdale’s s udies we e mainly based on he ba ium specimens a ailable a
di e en he ba ia. He himsel ecommended a comple e ield-based s udy o
esol e se e al species complexes in he genus.
Du ing he bo anical expedi ion o no heas e n India, we we e able o collec
a highly dis inc i olia e plan om A unachal P adesh, India. A e conduc ing
an ex ensi e li e a u e su ey pe aining o his genus (e.g. Lawson 1875; Cla ke
1881), we came ac oss Leea i olia a, which was i s desc ibed by Lawson in
1875 in Hooke ’s Flo a o B i ish India. Lawson (1875) desc ibed his species
as a small, wi y plan , 1–2 ee all and no ed ha i was allied o Leea aspe a
Wall. We we e able o loca e he syn ype ma e ial o Jenkins & c. (K000736385
& K000323729), Boo h (K000323730), and G i i h (K004252755, P02342863)
in K and P. Addi ionally, ou examina ion o he ex ensi e he ba ium specimens
om India (AHMA, ARUN, ASSAM, BSHC, BSI, BSID, CAL, CALI, MH, PBL, RHT
and TBGT) and he Uni ed S a es Na ional He ba ium (US), and o he collec-
ions i ually (A, BM, BR, G, GH, K, L, MICH, MO, NY, P, SING & U), enabled us o
loca e and compa e se e al his o ical collec ions o L. i olia a.
Leea i olia a was synonymized unde L. compac i lo a by Ridsdale (1974).
Ridsdale adop ed a b oade species concep o mos o he species. Leea i olia-
a is es ic ed only o India and Bhu an. Howe e , Leea compac i lo a, as desc ibed
by Ku z (1873) om Ma aban, Myanma , is a sh ubby species dis ibu ed in India,
Bhu an, China, Laos, Myanma , Nepal and Vie nam. Fo he ype ma e ial o L. com-
pac i lo a, we sea ched all possible he ba ia, especially he shee s deposi ed by
Ku z. Howe e , we we e unable o loca e a specimen by Ku z collec ed om he
ype locali y, Ma aban o e en om Myanma . In his e ision, Ridsdale (1974) was
also unable o loca e he specimen o Ku z. Hence, we need o ypi y he name.
The e o e, he aim o his pape is o examine he collec ions and mo pholo-
gy o bo h L. i olia a and L. compac i lo a o ee alua e he axonomic s a us
o L. i olia a and o neo ypi y he name L. compac i lo a.
Ma e ials and me hods
Ma e ials
The ma e ials o he p esen s udy we e ga he ed du ing he ield ip o no heas -
e n India. The axonomic key p o ided by Ridsdale (1974, 1976) and o he li e a-
u e dealing wi h his genus (Lawson 1875; Cla ke 1881; Gamble 1918; Nai hani
2000) as well as a la ge numbe o specimens we e s udied om a ious he ba ia
such as AHMA, ARUN, ASSAM, BSHC, BSI, BSID, CAL, CALI, MH, PBL, TBGT, A, BM,
BR, G, GH, K, KUN, L, MICH, MO, NY, P, SING, U and US (Ac onyms as pe Thie s
2024 con inuously upda ed) o assess he axonomic s a us o L. i olia a.
Mo phological analyses, dis ibu ion map, conse a ion assessmen s
and ypi ica ion
The desc ip ions we e p epa ed based on he li e ma e ials collec ed om
he ield and ma e ials plan ed in Calicu Uni e si y Bo anical Ga den, and by
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
examining he he ba ium specimens om ASSAM, ARUN and US he ba ia.
The specimens we e examined, and he pho omic og aphs we e aken us-
ing an M80 S e eomic oscope (Leica, Mannheim, Ge many) a ached wi h
a Leica MC 190 HD came a wi h Leica Applica ion Sui e Ve sion 4.12. To
ake he ield pho og aphs, Nikon D3400 and Nikon D750 came as (Nikon,
Japan) we e used. The desc ip ions we e p epa ed by using he e minolo-
gy o Ridsdale (1974, 1976), S ea n (1983), Ha is and Ha is (1995), Simp-
son (2019), and Kew Plan Glossa y (Been je 2016). A mic o- ip pen (Ro ing
a ian ) equipped wi h 0.1 and 0.2 poin s was used o he p epa a ion o
he illus a ion. The dis ibu ion map was p epa ed using he so wa e QGIS
e sion 3.24.3 (QGIS 2022). Du ing he ield isi s, he h ea was ho ough-
ly in es iga ed. Ex en o Occu ence (EOO) and A ea o Occupancy (AOO)
we e calcula ed using Geoca (h p://geoca .kew.o g/), and conse a ion
assessmen was pe o med using IUCN S anda ds and Pe i ions Commi -
ee (Ve sion 15.1 2022). Fo he selec ion o ypes, he mos sui able and
in o ma i e specimens we e chosen in concu ence wi h he p o ologue and
adhe ing o he p o isions o he Mad id Code (Tu land e al. 2025).
Resul s and discussion
Cla ke (1881) ecognized Leea i olia a and ea ed i in he se ies Rub i lo-
ae, which is cha ac e ized by ed-colo ed lowe s, in his e ision o Indian
Leea, bu admi ed he was no su e abou he colo o he co olla. Subse-
quen ly, Kanjilal and Das (1936) included his species in he Flo a o Assam.
Howe e , Ridsdale (1974) synonymized L. i olia a unde L. compac i lo a.
An ex ensi e ield explo a ion o bo h L. i olia a and L. compac i lo a, o-
ge he wi h he ba ium examina ions, suppo s he dis inc i eness o bo h
species. Leea i olia a di e s om L. compac i lo a by i s habi , lea a -
chi ec u e, pe iola s ipule, cha ac e is ics o pe iolule and achis, size o
in lo escence and shape and size o b ac s. Leea i olia a is a weak he -
baceous species, and he emaining species o he genus a e subsh ubs,
sh ubs, o small ees. Leea i olia a has a unique lea a chi ec u e wi h
mos ly i olia e o 1-pinna e lea es ha bea a basal pai o highly educed
lea le s (Figs 1, 2), consis en ly dis inc om hose o o he membe s o he
Leea. We hus eins a e L. i olia a and p o ide de ailed compa isons o he
wo species in Table 1.
Table 1. Mo phological compa isons o L. i olia a and L. compac i lo a.
Cha ac e s L. i olia a L. compac i lo a
Habi 0.8–1 (–2.5) m all, weak, he baceous, wi y plan O e 1 m s aggling o e ec sh ub
Lea a chi ec u e Mos ly i olia e o 1-pinna e, wi h a basal pai o lea le s
always much educed
Pinna e o bipinna e lea es
Indumen um on lea le s One ow o s i hai s be ween ne es on he uppe
su ace
Glab ous
Pe iolule and achis Pe iolules and achises winged Pe iolules and achises no winged
Pe iola s ipule Expanding he ull leng h o he pe iole, 2–2.5 cm long Hal o he leng h o he pe iole, 4–8 cm long
In lo escence size Up o 4 cm long 8–22 cm long
B ac s Lanceola e-linea B oadly o a e
Calyx Glab ous Hispid
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
Taxonomic ea men
Leea i olia a M.A.Lawson, Hook. . (Ed), Fl. B i . India 1(3): 666. 1875.
Figs 1, 2
Leea i olia a M.A.Lawson, Hook. . (Ed), Fl. B i . India 1(3): 666. 1875; C.B.
Cla ke, T imen’s J. Bo . 19: 163. 1881; Kanjilal & Das, Fl. Assam 1(2): 305.
1936. Type: INDIA. Assam, s.d., Jenkins & c. s.n. (lec o ype, designa ed by
Ridsdale 1974, pg. 88: K [K000736385]; isolec o ype: K [K000323729]).
Desc ip ion. Pe ennial weak he b, 0.8–1 (–2.5) m high; oo s non- ube ous.
S ems e ec o wi y, join ed, wa ed, glab ous; in e nodes u owed, g een;
nodes swollen; pea l glands globose, whi e. Lea es al e na e, i olia e o
1-pinna e wi h wo udimen a y pai s o lea es a base; pe ioles 7–21 mm
long, sligh ly bulged, shea hing he s em, hai y a base, g een. Pe iola s ipules
na owly winged, expanding he ull leng h o he pe iole, 2–2.5 cm long; s ip-
ula sca s iangula ; achises 1–4 cm long, channeled and sligh ly winged on
uppe su ace, hispid on ma gin, g een. Lea le s opposi e, ellip ic oblong o
obo a e, dis al lea le s lamina: 5.5–13 × 2.7–7.5 cm, la e al lea le s lamina:
4.5–12.5 × 2–6 cm, one ow o s i hai s in be ween ne es on uppe su ace,
ma u e lea es g een on uppe su ace and pale g een on lowe , base cunea e,
ma gins se a e, apex cuspida e cauda e, ab up ly acumina e; pe iolules: cen-
al 1–2.5 cm long, la e al 0.2–0.7 cm long, channeled and winged on uppe
su ace, g een; seconda y eins 7–16 pai s, 3-ne ed a base, s igose on lowe
su ace, ein aised on abaxial side. In lo escences e minal o lea -opposed,
compound cyme, up o 4 cm long, axis hispid. B ac s 1.8–2.7 mm long, lance-
ola e-linea , pubescen . Calyx campanula e, c. 3.5 mm long, 5 lobed, each lobe
c. 1–1.5 mm long, apex acu e; co olla lobes 5, 2.7–2.8 × 1.1–1.3 mm, sligh ly
hooded a apex, ma gin en i e. Flo al disc 1.1–1.5 mm long. O a y 4–6 locula .
F ui s no ed as black in Lawson (1875).
Flowe ing and ui ing. June–Decembe .
Habi a . This species mos ly g ows on oadsides and o es ma gins. They
g ow along wi h Adian um sp. (P e idaceae), Impa iens mino (DC.) Benne , Im-
pa iens balsamina L. (bo h Balsaminaceae) and some g asses.
Dis ibu ion. Bhu an, India (Assam & A unachal P adesh). Fig. 3.
Specimens examined. Bhu an. s.d., Boo h s.n. (K000323730). India. •
A unachal P adesh: Mishmi hills, s.d., G i i h 1344 (K004252755, P02342863);
• Eas Kameng dis ic , Pakke Wildli e Sanc ua y, 25 Ap 2012, B.B.T. Tham
35028 (ARUN); • Kamle dis ic , Raga, 27°47'59.4"N, 94°04'29.7"E, 16 Aug 2018,
Am u ha 159851(CALI); • Assam: s.d., Mas e s s.n (K004252752); • Cacha dis-
ic , Bhuban Hills, Oc 1978, 74142 (ASSAM); • Cacha , June 1874, R.L. Keenen
s.n. (K004252761); • Da ang dis ic , Aka Hills, s.d., N.L. Boh 17378 (ASSAM); •
Dib uga h dis ic , Dib u Rese e, eas e n pa , 20 Oc 1960, G. Panig ahi 21564
(ASSAM); • Golagha dis ic , s. loc, 300 , May 1891, D . King’s collec o s s.n.
(US01151426, K004252756, P02342865, L 0744660); • Jo ha dis ic , Gibbon
Sanc ua y, Camp-2, 12 June 2010, Ranji Daima y 122699 (ASSAM); • Ibid.,
Camp-4, 26 Feb 2011, Ranji Daima y 121691 (ASSAM); • Kama up dis ic ,
Moyang Residence, 09 Ap 1915, U. Kanjilal 5449 (ASSAM); Kama up Me o-
poli an dis ic , Guwaha i, Ap il 1902, A.C. Cha e jee (P02342864, LY0106862,
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
Figu e 1. Leea i olia a M.A.Lawson: A, D. Habi ; B. Adaxial su ace o lea le showing ichomes; C. Abaxial su ace o
lea le showing ichome on ne es; E, F. Pe iola s ipules; G. Rachis showing wings; H. S em; I–K. Lea es; L–N. In lo es-
cences wi h b ac s; Flowe Bud.
286
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
Figu e 2. A wig o Leea i olia a M.A.Lawson (D awn by Am u ha A. om Am u ha 159851).
287
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
MPU703118); • Ka bi Anglong dis ic , • Ga ampani Wildli e Sanc ua y, 24 No
1975, P.K. Haj a 65058 (ASSAM); Kaliani, 18 June 1963, D.B. Deb 35116 (AS-
SAM); • Nagaon dis ic , Laopani, 28 Aug 1978, P. Chak abo y 78984 (ASSAM);
• Soni pu dis ic , Ba asipu , 13 May 1947, M.M. S ini asan 22253 (ASSAM); •
Kanchanju i, Kazhi anga, 12 Aug 1973, P.K. Haj a 52836 (ASSAM); • Tinsukia
dis ic , Ma ghe i a Fo es , 22 June 1938, G.K. Duka 17010 (ASSAM).
Conse a ion s a us. Leea i olia a is es ic ed o wo no heas e n Indian
s a es, Assam and A unachal P adesh, and one loca ion in Bhu an. This spe-
cies p e e s o g ow in open a eas adjoining e e g een o es s. The Ex en o
Occu ence (EOO) is calcula ed o be 114,777 km2, and he A ea o Occupan-
cy (AOO) is 84 km2. Conside ing i s la ge Ex en o Occu ence, his species
does no all in o any h ea ca ego y. Despi e his, he A ea o Occupancy is
small, and each popula ion has ewe han i e ma u e indi iduals. Due o i s
na ow ange o dis ibu ions and small AOO, his species could be ca ego ized
a p esen as Endange ed (EN), B2b(ii,iii) acco ding o he IUCN S anda ds and
Pe i ions Commi ee (2022).
No es. In he p o ologue, Lawson (1875) desc ibed he an he s as dis inc ,
and he ui s we e black.
Ridsdale (1974) ypi ied he name L. i olia a by selec ing he Jenkins & c.
ma e ial om Assam as he lec o ype. We we e able o ace wo shee s o Jen-
kins & c. om he Kew he ba ium (K000736385 & K000323729). Howe e , Rids-
dale has a ached a slip o he shee wi h ba code K000323729 s a ing ha i is
he lec o ype. Hence, he o he shee (K000736385) becomes he isolec o ype.
Figu e 3. Dis ibu ion o Leea i olia a M.A.Lawson in no heas e n India and Bhu an (d awn using QGIS e sion 3.24.3).
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
Leea compac i lo a Ku z, J. Asia . Soc. Bengal, Na . His . 42(2): 65. 1873
Figs 4, 5
Leea compac i lo a Ku z, J. Asia . Soc. Bengal, Na . His . 42(2): 65. 1873, Fo es
Fl. Bu ma 1: 279. 1877; C.B.Cla ke in T imen’s J. Bo . 19: 163. 1881; Ridsdale,
Blumea 22: 88. 1974; J.Joseph, Fl. Nongpoh and icini y 72. 1982; Ha id. &
R.R.Rao, Fo es Fl. Meghalaya 248. 1985; A.S.Chauhan, Con . Fl. Namdapha
140. 1996; Haj a e al., Ma . Fl. A unachal P adesh 1: 318. 1996; B.D.Nai hani
in N.P.Singh e al. (Ed), Fl. India 2000; R.Shanp u in N.P.Singh e al. (Eds), Fl.
Manipu 1: 240. 2000; N.P.Singh e al., Fl. Biha , Analysis 114. 2001; T.K.Sa ma
& A.K.Sa ka , Fl. Palamau Dis . 156. 2002; K.P.Singh in N.P.Singh e al. (Eds),
Fl. Mizo am 1: 388. 2002; G.D.Pal, Fl. Lowe Subansi i Dis . 1: 215. 2103, excl.
syn. L. i olia a; B.K.Shukla in K.P.Singh e al. (Eds), Fl. U a P adesh 1. 351.
2016, excl. syn. L. i olia a; S.S.Dash & P.Singh, Fl. Ku ung Kumey Dis . 384.
2017, excl. syn. L. i olia a; Pusalka & S.K.S i as ., Fl. U a akhand 1: 989.
2018. Type: INDIA. Nagaland: Naga hills, 3 No 1836, G i i h He b. P op . N.
1297 (neo ype, K K004252754, he e designa ed).
Leea b ac ea a C.B.Cla ke, T imen’s J. Bo . 19: 164. 1881; Kanjilal & Das, Fl.
Assam 1(2): 308. 1936; Deb, Fl. T ipu a 1: 419. 1981; A.K.Mukh., Pachma hi
& Bo i Rese es 68. 1984. Type: INDIA. Sikkim, s.d., Hook. . & Thomson 169
(lec o ype, designa ed by Ridsdale 1974, pg. 88: K [K001615004]).
Desc ip ion. E ec subsh ub o sh ub, 1–2 m high; oo s non- ube ous.
S ems e ec o s aggling, s ou , join ed, wi h b anches, young s em hispid
o pubescen ; in e nodes u owed o ound, g een; nodes no swollen; pea l
glands globose, mo e on in lo escences, whi e. Lea es al e na e, pinna e o
bipinna e, impa ipinna e; pe ioles 7–21 mm long, base bulged, shea hing
he s em, glab ous, g een. Pe iola s ipules na owly winged, 4–8 cm long,
ex ending o hal o pe ioles; s ipula sca s iangula ; achises 5–31 cm
long, channeled on uppe su ace, hispid, g een. Lea le s opposi e, o a e
widely o a e o obo a e, dis al lea le s: lamina 14–32 × 6.5–13.5 cm, la e al
lea le s: lamina 7.5–30 × 4–13 cm, glab ous, young lea es g een o b own,
ma u e lea es g een on uppe su ace and pale g een on lowe , base cunea e
o ounded, ma gins se a e, apex long acumina e o cauda e; pe iolules:
cen al 3–8 cm long, g een, la e al 0.4–2.5 cm long; seconda y eins 7–16
pai s, 3-ne ed a base, ein aised on abaxial side. In lo escences e minal,
a lea -opposed compound cyme, 8–22 cm long, many lowe ed, condensed,
hispid; peduncles 1–4 cm long, u owed, pubescen , g een, hispid. Flow-
e s 4–6.2 mm, ac inomo phic, g eenish wi h b own inges. Pedicels slende ,
1.5–2 mm long, hispid. B ac s 3.6–8.3 × 1.9–5 mm, b oadly o a e, conspic-
uous, pe sis en , g een, hispid. Calyx campanula e, 3.5–5.6 × 3–3.3 mm,
5 lobed, lobes 0.9–1.1 × 0.75–1.9 mm, o a e o widely o a e, apex acu e,
g een, hispid; co olla lobes 5, o a e-oblong, 2.7–3.5 × 1.9–2.4 mm, apex
acu e, hooded, ma gin en i e, glab ous, g een; co olla ube wi h lo al disc
3.1–3.3 × 3–3.2 mm. Flo al disc 1.6–2.4 mm long, o ange- ed; uppe ee
pa 1.1–1.8 mm long, no ched; sinuses shallow, 0.25–0.4 mm long; lowe
ee pa 0.8–1 mm long. S amens 5, syngenecious; ilamen s 1.25–1.3 mm
long, glab ous, whi e; an he s 1.2–1.3 mm long, dehiscing longi udinally,
pu ple on backside, c eamish on ma gin. O a y globose, 0.95–1.1 mm long,
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Am u ha A halappil e al.: Resol ing he axonomy o Leea i olia a
Figu e 4. Leea compac i lo a Ku z: A. Habi ; B. F ui ing wig; C. Pe iola s ipule; D1, D2. Lea le s; E. In lo escence; F. In-
uc escence; G. B ac ; H. Flowe bud; I. Flowe ; J. Calyx; K. Co olla lobe; L. Co olla ube wi h lo al disc; M. Co olla ube
wi h lo al disc spli open; N. S amens; O. Pis il; P. Seeds.
