Pinguicula pan e iae (Len ibula iaceae), a new
Ca ibbean bu e wo om eas e n Cuba
I an Pančo1, Paul Temple2, Geo anys Rod íguez Cobas3, Noel Cou in Lobaina3,
I ana Ma iso a4, Yoannis Domínguez5
1 Ni ica, Slo akia
2 Cons anza, Dominican Republic
3 Unidad de Se icios Ambien ales Alejand o de Humbold , Ba acoa, Guan ánamo, Cuba
4 Po ubka, Slo akia
5 Uni e sidade Es adual Paulis a (UNESP), Ins i u o de Biociências, São Vicen e, SP, B azil
Co esponding au ho s: I an Pančo (i an.pan[email p o ec ed].uk), Yoannis Domínguez ([email protected] )
Academic edi o : And é Simões ♦
Recei ed
14 June 2025 ♦
Accep ed
20 Augus 2025 ♦
Published
8 Oc obe 2025
Abs ac
Backg ound and aims – Pinguicula is a cosmopoli an genus o ca ni o ous plan s wi h mo e han 100 species, mos o
hem dis ibu ed in wo cen es o di e si y: Eu ope and he Cen al Ame ican-Ca ibbean egion. Fou een species a e
epo ed o Cuba. Du ing ieldwo k o s udy Pinguicula popula ions in eas e n Cuba, a whi e- lowe ed axon, p e iously
misiden i ied as Pinguicula albida o Pinguicula ja aguana, was collec ed in he moun ainous egions o he Alejand o de
Humbold Na ional Pa k. This axon ep esen s a new species o which we p esen a ull desc ip ion, a dis ibu ion map,
illus a ions, pho og aphs, and a conse a ion s a us.
Ma e ial and me hods – Li ing and p ese ed ma e ial o se e al popula ions o whi e- lowe ed Pinguicula om Cuba
we e analysed, measu ed, and desc ibed based on ege a i e and ep oduc i e cha ac e s. S a is ical analyses we e ca ied
ou o explo e he di e ences among he collec ions. The p elimina y conse a ion s a us o he new species was assessed
based on he IUCN guidelines and c i e ia.
Key esul s – Pinguicula pan e iae is desc ibed as a new species endemic o eas e n Cuba. I h i es on pe manen ly
we , e i ic soils de i ed om se pen ine bed ocks, on slopes abo e he banks o he Ri e Toa in associa ion wi h
su ounding semi-a id mon ane se pen ine sh ubwood (cha ascal). I di e s om he closely ela ed species, Pinguicula
albida and Pinguicula ja aguana, p ima ily by i s smalle ose es wi h obo a e, sube ec o e ec lea es, non-in olu e lea
ma gins, smalle lowe s up o 1–1.3 cm including he spu , longe (0.6–0.7 mm) and wide (0.18–0.22 mm) seeds, and
ha i g ows in ull sun.
Conclusion – The newly desc ibed species inc eases he numbe o Pinguicula in he Cuban a chipelago o 15. Pinguicula
pan e iae is p elimina ily assessed as C i ically Endange ed (CR) acco ding o he IUCN Red Lis Ca ego ies and C i e ia.
Keywo ds
bu e wo , ca ni o ous plan , c i ically endange ed, endemic, Pinguicula albida, Pinguicula ja aguana, axonomy
INTRODUCTION
Cuba is a ema kably ich sou ce o di e si y o he
genus Pinguicula L., conside ing i is an island wi h an
a ea o only 110,860 km2. The dis ibu ion o he genus
Pinguicula in Cuba shows a disjunc ion o e h ee egions
– wes , cen al, and eas – wi h signi ican gaps be ween
he egions. O he 14 Cuban species cu en ly accep ed
(Pan e 2024), nine occu wi hin he eas e n egion.
Be o e he middle o la e 1960s, only i e species had
been eco ded o he en i e coun y: Pinguicula albida
C.W igh ex G iseb., P. benedic a Ba nha , P. ili olia
C.W igh ex G iseb., P. jackii Ba nha , and P. lignicola
Ba nha (Alain 1957; E ns 1961; Caspe 1966). This
Plan Ecology and E olu ion 158 (3): 382–391, 2025
h ps://doi.o g/10.5091/plece o.161641
Copy igh I an Pančo, Paul Temple, Geo anys Rod íguez Cobas, Noel Cou in Lobaina, I ana Ma iso a, Yoannis Domínguez. This is an open access a icle
dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any
medium, p o ided he o iginal au ho and sou ce a e c edi ed.
Plan Ecology and E olu ion is published by Meise Bo anic Ga den and he Royal Bo anical Socie y o Belgium.
RESEARCH ARTICLE
Plan Ecology and E olu ion 158 (3): 382–391, 2025 383
changed om 1966 onwa ds when bo anical expedi ions
we e o ganised in Cuba as pa o he wo k, Flo a de la
República de Cuba (Méndez 1995; Regalado Gabancho
e al. 2008), which yielded nume ous obse a ions and
collec ions o Pinguicula, including se e al new species.
The esul ing specimens a e kep a he He ba ium P o .
Johannes Bisse (HAJB, Na ional Bo anic Ga den o
Cuba, Uni e si y o Ha ana) wi h duplica es in Ge man
he ba ia (B, Bo anic Ga den and Bo anical Museum
Be lin-Dahlem and JE, F ied ich-Schille -Uni e si ä
He ba ium Haussknech ).
Du ing his pe iod o bo anical ield ips, he iden i ies
o some specimens o whi e- lowe ing Pinguicula om
he eas e n egion o Cuba emained unde e mined
(HFC-21959a, Ebel s.n. [HAL 36587], HFC-39913) bu
se e al we e mis akenly iden i ied as P. albida (HFC-
16747, HFC-16918, HFC-21649a, HFC-21649b, HFC-
21959b, HFC-21959c, HFC-56240b, HFC-56240c). This
axon was p e iously and co ec ly only known om he
wes o Cuba. A ha ime, only h ee whi e- lowe ed
axa we e known, P. albida, he epiphy ic P. lignicola, and
some popula ions o P. ili olia, he la e wo could no
be con used wi h o he Cuban Pinguicula species because
o hei dis inc ly unique mo phological di e ences.
Likewise, eas e n Pinguicula wi h non-whi e lowe s we e
commonly de e mined o be P. benedic a (HFC-56240a)
based on supe icial simila i y.
Bisse e al. (1975) discussed an eas e n Pinguicula
iden i ied as P. albida (HFC-16747) and ou lined se e al
di e ences be ween hese plan s, collec ed a Cayo
Fo una (loca ed on he Ri e Toa), and he mo e ypically
wes e n popula ions bu dismissed hei impo ance. Thus,
despi e an ea ly indica ion by F. Ebel in 1972 ha ano he
specimen om Cayo Fo una (Ebel s.n. [HAL 37587], in
sched.) was a new species, he iew was main ained ha P.
albida was no es ic ed o wes e n Cuba.
Fo una ely, he la e Sieg ied Jos Caspe ecognised
ha many o he he ba ium specimens om eas e n Cuba
we e new axa. His desc ip ions (Caspe 2003, 2004, 2007,
2019) allowed a be e compa ison be ween he species,
especially hose om he eas ha we e signi ican ly less
well known. Among hese new species was he whi e-
lowe ed Pinguicula ja aguana Caspe (Caspe 2003),
o which he ype ma e ial (HFC-39913) was collec ed
beside a s eam, he A oyo Ja agua, wi hin he Alejand o
de Humbold Na ional Pa k, whe e i is gene ally ound
g owing on e ical we walls in shaded loca ions. As a
esul o sepa a ing eas e n whi e- lowe ed species in o P.
ja aguana, he ea e , P. albida was conside ed endemic o
wes e n Cuba and all whi e lowe ing species om eas e n
Cuba we e conside ed o be P. ja aguana. Howe e ,
Caspe was unawa e o a numbe o specimens om
sepa a e ga he ings om Cayo Fo una, a egion wi hin
he Oji o de Agua sec o o he same pa k. The e o e,
hese specimens emained unci ed and wi hou p ope
de e mina ion. La e , Caspe (2019) published a e ision
o he genus in he G ea e An illes and included he Cayo
Fo una specimens wi hin his concep o P. ja aguana.
He no ed ha he e a e di e ences in lowe mo phology
be ween he plan s om he P. ja aguana locus classicus
beside he A oyo Ja agua and hose om Cayo Fo una.
Despi e hese di e ences, he concluded ha he
obse ed di e ences we e wi hin he ange o a ia ion
o P. ja aguana. Thus, all Cayo Fo una collec ions we e
subsumed unde P. ja aguana. By 2007, he ea ly F. Ebel
ga he ing (Ebel s.n. [HAL 36587]), o iginally de e mined
as “Pinguicula no . spec.”, had cap u ed Caspe ’s a en ion
and he newly de e mined i as “Pinguicula c. . ja aguana”.
La e s ill, his specimen, as well as a colou pho og aph
o li ing plan s in habi a also supplied by F. Ebel, was
ci ed by Caspe (2019); his explici men ion o he Cayo
Fo una popula ion, whe e he used hose plan s as a
compa ison o P. ja aguana wi h P. albida, demons a ed
his inclusion o he popula ion wi hin P. ja aguana. By
his ime, o he specimens om Cayo Fo una we e also
subsumed wi hin P. ja aguana, likely because hey lacked
lowe s (HFC-16747 and HFC-16981). Howe e , ou
p elimina y analyses o he ba ium specimens e ealed
ha , wi hou excep ion, specimens om his locali y do
no ep esen he P. ja aguana species concep .
In ecen yea s (2019–2025), se e al expedi ions
had been unde aken o s udy popula ions o eas e n
Cuba Pinguicula. Mo phological analyses and
ield obse a ions allowed he ecogni ion o clea
mo phological and ecological di e ences be ween he
popula ion o Pinguicula om Cayo Fo una and all
o he known whi e- lowe ed species hese plan s ha e
been con used wi h. Thus, a new species o Pinguicula
om eas e n Cuba is he ein desc ibed and illus a ed. I s
axonomic ela ionships, dis ibu ion and ecology, and i s
p elimina y conse a ion s a us a e also p esen ed.
MATERIAL AND METHODS
Field ips we e conduc ed be ween No embe 2019
and Ma ch 2025 o s udy Pinguicula popula ions in
eas e n Cuba. Plan s om a single popula ion p e iously
a ibu ed o P. ja aguana, loca ed a Cayo Fo una, (a
sec ion o he Ri e Toa), we e examined and compa ed
wi h ype ma e ial and specimens (Supplemen a y
ma e ial 1) o Pinguicula albida and P. ja aguana om
he ba ium collec ions (A, B, BRU, GH, GOET, HAC,
HAJB, HAL, HFA, HPPR, HUMC, JE, MA, MO, MT, NY,
US, and YU; ac onyms ollow Thie s 2025). Collec ed
ma e ial was also compa ed o desc ip ions in he ele an
li e a u e (Alain 1957; E ns 1961; Caspe 1966, 2003,
2004, 2007, 2019; Bisse e al. 1975; Domínguez e al. 2014;
Temple e al. 2023).
Mac o- and mic omo phological analyses o ecen ly
collec ed specimens and olde specimens in he ba ium
collec ions, as well as ield obse a ions, we e ca ied
ou o desc ibe he new axon. Field pho og aphs we e
aken using a Canon R6 Ma k 2 came a wi h 100 mm 2.8
mac o (Canon, Japan) o Laowa 25mm 2.8 2.5–5× ul a
mac o lens (Venus Op ics, China). Mic omo phological
cha ac e s we e analysed and pho og aphed using a BEL
Pančo e al.: Pinguicula pan e iae (Len ibula iaceae), a new species om Cuba384
mic oscope model SOLARIS-T wi h a coupled digi al
came a BEL EU 12 CONVS (Bel Pho onics®, China) using
he so wa e Cap u e 2.1 (Meiji Techno, Japan). Eigh
quan i a i e ege a i e and ep oduc i e cha ac e s (Table
1; Supplemen a y ma e ial 2) we e eco ded o each
axon (n = 60). The no mali y o he da a was assessed o
each a iable indi idually using he Shapi o-Wilk’s es
(p alues = 0.000–0.020; Supplemen a y ma e ial 3). To
compa e mo phome ic da a, a mul i a ia e analysis was
pe o med o educe dimensionali y and ob ain new axes
ha be e explain da a a ia ion. A p incipal componen
analysis (PCA) based on he co ela ion ma ix was
pe o med, wi h he absolu e alues ans o med o
z-sco es and plo ed in a wo-dimensional space de ined
by he i s wo p incipal componen s (Fac o 1 and 2).
All s a is ical analyses we e pe o med using SYSTAT 13
.13.2.01 (Sys a So wa e, Inc. 2023).
The p elimina y conse a ion s a us o he new species
was assessed ollowing IUCN Red Lis Ca ego ies and
C i e ia (IUCN 2024), o which he ex en o occu ence
(EOO) and he a ea o occupancy (AOO, wi h 2 × 2
km g id) we e calcula ed using he so wa e GeoCAT
(Bachman e al. 2011). The dis ibu ion map, based on he
known dis ibu ion o he species acco ding o he ba ium
eco ds and ield su eys, was p epa ed wi h QGIS
.3.40.3 (QGIS De elopmen Team 2024).
TAXONOMIC TREATMENT
Pinguicula pan e iae Panco, P.Temple & Y.Domínguez,
sp. no .
u n:lsid:ipni.o g:names:77357113-1
Figs 1–4A
Type. CUBA – Guan ánamo • Ya e as, Pa que Nacional
Alejand o de Humbold , Palenque, Cayo Fo una, cuenca
del ío Toa, 3 km desde la es ación Pied a La Vela;
20°24’54.6”N, 74°57’39.5”W; 460 m; 31 Ma . 2023; J.
Imbe Planche, I. Pančo, G. Rod íguez Cobas, I. Ma iso a,
N. Cou in Lobaina s.n.; holo ype: HAJB [G 002601];
iso ypes: B, BSC [22092].
Diagnosis. Pinguicula ja aguana Caspe a inis sed
b e ibus, oliis o oba is, e ec is sube ec is, ma gine
in ege ima non in olu e, lo es pa i 10–13 mm (calca i
incluso), semina 0.6–0.7 mm longa (appendix incluso),
0.18–0.22 mm la a.
Desc ip ion. Pe ennial he b, scapose, in a basal ose e,
(19–)25–30(–35) mm in diame e ; sube ec o e ec ,
some imes g owing densely g ouped oge he in colonies
o many indi iduals exhibi ing cala hi o m ose es.
Rhizome sho , wi h ib ous oo s. Lea es (4–)5–7(–10),
homophyllous, yellow-g een o eddish-b own, (10–)12–
15(–17) × (6–)8–10(–11) mm, obo a e, apex ounded,
base na owly cunea e in o a sho pe iole, ma gins
en i e; adaxial su ace densely co e ed wi h glandula
ichomes. Scapes 1(–3), 1- lowe ed, e ec , g een o
eddish-b own, (50–)80–110(–150) mm long, ± 0.30–
0.45 mm hick, spa sely glandula wi h e y ew s alked
and sessile ichomes. Flowe s whi e, indis inc ly 2-lipped
(subisoloba e). Calyx dis inc ly 2-lipped, simila colou
o lea es, sho e han he ube, ± 4 mm; ou e su ace
spa sely co e ed wi h s alked glandula ichomes; uppe -
lip di ided nea ly o he base in o 3 lobes, lobes oblong-
ob use, apex e use, ± 2 × 1 mm; lowe lip 2-lobed, he
lobes ee nea ly om hei bases. Co olla (8.5–)9–10(–
11.3) mm long (including he spu ), ± 9 mm diame e ,
widely opened; he lobes o ming an angle wi h he ube o
± 90°, b oadly obo a e, ma gins o e lapping o less han
hal hei leng h, whi e, (3.8–)4–4.7(–5.2) × (2.9–)3.5–
3.8(–4.3) mm, adaxial su ace co e ed spa sely wi h
non-glandula ichomes ha a e mo e abundan a he
en ance o he ube; uppe lip 2-lobed, lobes smalle han
he wo la e al lowe -lip lobes; lowe -lip deeply 3-lobed,
lobes obo a e, apex ounded, midlobe (4.5–)4.8–5(–5.2)
× (3.8–)4(–4.2) mm. Tube yellow wi h deep b own eins,
cylind ic, sho , (2.5–)3–4(–4.5) × 1.5–2.5 mm, wi hou
pala e, dis inc ly ben (genicula e) and held a an angle
o ± 90°, ipa i e, he en ance wi h long unise ia e
non-glandula ichomes, he middle pa wi h medium-
leng h unise ia e and s ipi a e glandula ichomes,
he inne pa wi h sho glandula ichomes. Spu
yellowish-b own, hick, conical, e y sho , (2.5–)3–4(–
4.5) × 1.5–2.5 mm, apex ounded, sacca e. S amens ± 1.5
mm long, sligh ly cu ed; an he s ± 0.5 mm wide. Pollen
p ola e (P:E = 1.49), zonoape u a e, 4(5)-colpo a e,
(34.63–)35.52–37.32(–39.17) × (23.53–)24.12–24.88(–
Table 1. Quan i a i e ege a i e and ep oduc i e cha ac e s used in he s a is ical analyses, codes, and desc ip ions. All
measu emen s we e aken in mm.
Cha ac e Code Desc ip ion
1 Lea leng h LFL Measu ed om he base o he lea blade o i s apex
2 Lea wid h LFW Measu ed a he wides pa o he lea blade
3 Scape leng h SCL Measu ed om he cen e o he ose e o he apex o he in lo escence
4 Flowe leng h FLL Measu ed om he ip o he spu o he apex o he co olla midlobe
5 Co olla lobe leng h (excep he midlobe) LBL Measu ed om he apex o he co olla ube o he apex o he lobe
6 Co olla lobe wid h (excep he midlobe) LBW Measu ed a he wides pa o he lobe
7 Co olla midlobe leng h MLL Measu ed om he apex o he co olla ube o he apex o he midlobe
8 Co olla midlobe wid h MLW Measu ed a he wides pa o he midlobe
Plan Ecology and E olu ion 158 (3): 382–391, 2025 385
Figu e 1. Pinguicula pan e iae based on ype specimens (holo ype HAJB [G 002601], iso ypes B, BSC [22092]) and on li ing ma e ial.
A. Habi , pe ennial he b wi h de ails o he ichomes o he lea and he in lo escence scape (ci cles). B. Eme ging lea wi h in olu e
ma gin. C. F on al iew o he lowe . D. La e al iew o he lowe . E1. In e nal ichomes o he ou e pa o he co olla ube. E2.
In e nal ichomes o he middle pa o he co olla ube. E3. In e nal ichomes o he inne pa o he co olla ube. F. Dissec ed
lowe wi h he pe sis en calyx and la e al iew o he s amens and gynoecium. G. F ui wi h pe sis en calyx. H. La e al iew o he
seed. Illus a ion by Yoannis Domínguez.
Pančo e al.: Pinguicula pan e iae (Len ibula iaceae), a new species om Cuba386
Figu e 2. Pinguicula pan e iae a he ype locali y in he Alejand o de Humbold Na ional Pa k, eas e n Cuba. A. Habi . B. F on al
iew o he lowe . C. La e al iew o he lowe . D. Dissec ed lowe showing he pe sis en calyx and s amens and gynoecium. E.
La e al iew o he ui . F. Habi a a he banks o he Ri e Toa. G. Regula ose es wi h sube ec lea es. H. Cala hi o m ose es
wi h e ec lea es. Pho os by I an Pančo (A–E, G–H) and I ana Ma iso a (F).
Plan Ecology and E olu ion 158 (3): 382–391, 2025 387
25.39) μm. O a y subglobose-o oid, sessile, ± 1.5 mm
in diame e , 1-locula e, ending in a sho s yle, glab ous.
S igma sho , 2-lobed, ma gin inged. Capsule globose, ±
2 × 2 mm. Seeds da k b own, usi o m, (440–)520–540(–
590) × (180–)200–215(–220) μm excluding appendage,
es a e icula e, myc opila appendage p ominen ,
(60–)80(–90) μm.
E ymology. Pinguicula pan e iae is named in honou o
he la e Cuban bo anis , P o . D C is ina Me cedes Pan e
Valdés, o acknowledge he con ibu ions o Cuban lo a
esea ch, in pa icula , wi h ega d o he ca ni o ous
plan amilies o which she was he main Cuban
specialis . Besides he esea ch, P o . Pan e dedica ed
mos o he li e o empowe ing many gene a ions o
Cuban bo anis s o achie e p o essional s anding, which
is also acknowledged he ein.
Phenology. Based on he p esence o lowe s o ui s,
o bo h, in he analysed specimens and acco ding o he
in si u obse a ions, he lowe ing pe iod ex ends om
Feb ua y o June.
Dis ibu ion and habi a . Pinguicula pan e iae is endemic
o eas e n Cuba (Fig. 3) whe e i is limi ed o a small a ea
wi hin Cayo Fo una, a subdepa men o he Oji o de
Agua sec o o he Alejand o de Humbold Na ional Pa k.
Plan s a e ound a an al i ude o ca 460 m, on s eep slopes
abo e he banks o he Ri e Toa (Fig. 2F), whe e hey
g ow in dense pa ches o alling a ew housand indi iduals.
Whe e a g oup o plan s g ow in a igh a angemen ,
hey ypically exhibi cala hi o m (cup-like) ose es wi h
sube ec o e ec lea es (Fig. 2H). While we can be ce ain
o he loca ion o Pinguicula pan e iae in Cayo Fo una,
he eco ds o he loca ions o o he specimens men ion
Palenque, Cayo Fo una, o eas e n Cuba (O ien e) bu
he use o such names is no de ailed enough o de e mine
whe he any co esponds o he ue Cayo Fo una o o
one o mo e o he loca ions. The ege a ion a he ype
locali y is semi-a id mon ane se pen ine sh ubwood
(cha ascal) ha occu s in associa ion wi h e i ic
soils de i ed om se pen ine bed ocks (Bo hidi 1996).
Howe e , Pinguicula pan e iae g ows on pe manen ly
we landslides and seeps on o jus abo e i e s banks.
Despi e he semi-a id condi ions o he cha ascal, his
a ea suppo s edaphic climax communi ies de eloped
unde a humid opical ain o es clima e.
Rela ionships. Acco ding o he in agene ic classi ica ion
p oposed by Fleischmann (2021), Pinguicula pan e iae
belongs o subg. Temnoce as Ba nha , sec . Homophyllum
Caspe , which also includes all Ca ibbean species om
Cuba and he Dominican Republic. Among he Cuban
species, Pinguicula pan e iae shows close mo phological
a ini ies wi h P. albida and P. ja aguana as hey ha e whi e
lowe s wi h subisoloba e co olla and a hick, sho spu .
Howe e , mo phological and mo phome ic analyses
ha e shown ha hese h ee species can be sepa a ed on
he basis o ege a i e and ep oduc i e ai s (Figs 4, 5).
In addi ion o he di e ences highligh ed in he diagnosis,
a compa a i e o e iew o Pinguicula pan e iae, P. albida,
and P. ja aguana is p o ided in Table 2.
P elimina y IUCN assessmen . Ve y ew eco ds exis
o his species acco ding o he analysed ma e ial in
he ba ium collec ions and, due o he limi ed access o
ma e ial and insu icien in o ma ion associa ed wi h
old specimens (1970–1972), i is di icul o de e mine i
all o hem o igina ed om a single popula ion. Caspe
(2019) ema ked ha se e al o he independen sigh ings
o plan s, he ein desc ibed as Pinguicula pan e iae, may
all ep esen eco ds o he same and cu en ly only
known popula ion (Fig. 3). Obse a ions om ou ield
explo a ions in eas e n Cuba suppo Caspe ’s iew. This
popula ion al eady has he highes Cuban p o ec ion
s a us and comp ises housands o plan s. Ne e heless,
he popula ion is es ic ed o small pa ches in a
Figu e 3. Dis ibu ion o Pinguicula pan e iae, Pinguicula albida, and Pinguicula ja aguana in Cuba based on he ba ium eco ds and
ield su eys.
Pančo e al.: Pinguicula pan e iae (Len ibula iaceae), a new species om Cuba388
Table 2. Compa ison o mo phological and habi a di e ences be ween Pinguicula pan e iae, P. albida, and P. ja aguana. Diagnos ic
cha ac e s o P. pan e iae a e indica ed by an as e isk (*).
P. pan e iae P. albida P. ja aguana
Rose e diame e *
(mm) (19–)25–30(–35) (28–)34–50(60) (19–)28–42(–60)
Lea
Type Homophyllous, wi h consis en ly
uni o m ca ni o ous lea es
Homophyllous, wi h consis en ly
uni o m ca ni o ous lea es
Homophyllous, wi h ca ni o ous
lea es exhibi ing seasonal
a ia ion o shape and size
G ow h* Sube ec o e ec Fla , adp essed o he subs a e Fla , adp essed o he subs a e
Size* (mm) (10–)12–15(–17) × (6–)8–10(–11) (14–)17–20(–45) × (9–)10–20(–30) (10–)19–20(–31) ×
(4–)8–8.5(–13)
Shape* Obo a e O a e o subo bicula e Oblong o spa hula e
Ma gin* En i e, non-in olu e In olu e In olu e
Flowe
Lobe size* (mm) (3.8–)4–4.7(–5.2) ×
(2.9–)3.5–3.8(–4.3) (5–)7–8(–10) × (4–)5–7(–9) (6–)9–10(–13) × (4–)6–7(–12)
Tube (mm) (2.5–)3–4(–4.5) × 1.5–2.5 (1.5–)2–3.5(–4.5) ×
(1.8–)2.5–3.2(–4) (2.5–)3–4(–5) × 1.5–2.5
Spu (mm) (2.5–)3–4(–4.5) × 1.5–2.5 (1.9–)3.3–4.6 × (0.7–)1–1.4(–1.8) (3–)4–5(–6) × (2–)2.5–3(–6)
Spu - ube angle Righ Righ Acu e
Capsule
Size (mm) 1.9–2.1 × 1.9–2.1 1.8–2.5 × 1.5–2.3 3.3–3.9 × 3.3–3.9
Seed
Size (including
appendage)* (μm)
(595–)610–625(–670) ×
(180–)200–215(–220)
(304–)418–480(–540) ×
(120–)150–170(–201)
(430–)500–580(–650) ×
(130–)150–200(–210)
Habi a In ull sun, in semi-a id mon ane
se pen ine sh ubwood (cha ascal)
In shade, in g assland, pine sa anna
and pine-mixed sh ubwood, d y
lowland sh ubwood (cuabal)
In shade, in semi-a id mon ane
se pen ine sh ubwood
(cha ascal)
Figu e 4. Dis inc ege a i e and ep oduc i e mo phological ai s o ela ed species. A. Pinguicula pan e iae. B. Pinguicula albida.
C. Pinguicula ja aguana. Pho os by I an Pančo (A, C) and Yoannis Domínguez (B).
Plan Ecology and E olu ion 158 (3): 382–391, 2025 389
Table 3. Loading o he i s wo axes (Fac o 1 and 2) o he p incipal componen analysis on quan i a i e mo phological a iables.
Cha ac e codes co espond o hose in Table 1.
Cha ac e Fac o 1 Fac o 2
LFL 0.306 0.842
LFW -0.458 0.774
SCL 0.782 -0.255
FLL 0.958 -0.026
LBL 0.938 0.073
LBW 0.941 0.174
MLL 0.968 0.096
MLW 0.957 -0.005
Va iance explained by componen s 5.451 1.419
Figu e 5. P incipal componen analysis based on ege a i e and ep oduc i e quan i a i e ai s o Pinguicula pan e iae, Pinguicula
albida and Pinguicula ja aguana. A. P ojec ion o he analysed a iables on o he i s wo p incipal componen axes (Fac o 1 and
2). B. P ojec ion o he sampled indi iduals o each species on o he i s wo p incipal componen axes (Fac o 1 and 2). Cha ac e
codes co espond o hose in Table 1.
na owly de ined a ea; mos occu on dis u bed g ound
om ecen landslides whe e hey a e a isk o u he
simila e en s. The e o e, based on he numbe o known
loca ions (1), he ex en o occu ence and he a ea o
occupancy (EOO/AOO = 0.810 km2), he p elimina y
conse a ion s a us p oposed is C i ically Endange ed
CR B1ab(i,ii,i )+B2ab(i,ii,i ) ollowing IUCN c i e ia
(IUCN 2024).
Addi ional ma e ial s udied (pa a ypes). CUBA –
O ien e [Guan ánamo] • Palenque, Cuchillas de Toa,
Cayo Fo una, a lo la go del ío Toa; Ap . 1970; J. Bisse
HFC-16747; HAJB • Palenque, Cuchillas de Toa, Cayo
Fo una, a lo la go del ío Toa; Ap . 1970, J. Bisse HFC-
16918; HAJB • 35 km no dös lich on Guan anamo,
Gebie on Palenque, Cayo Fo una, Au übe iesel em
Felsen am U e des Rio Toa; 22 Ma . 1972; F. Ebel s.n.;
HAL [36587].
DISCUSSION
Mo phological di e ences obse ed among he h ee
species a e use ul o cha ac e ise and iden i y hem
(Fig. 4; Table 2) and showed consis ency among he
analysed indi iduals o each axon. Fu he mo e, he
esul s o he PCA e ealed wo p incipal componen s
ha oge he explain 81.86% o he o al a ia ion in he
da a (Fac o 1: 68.13%, eigen alue 5.451; Fac o 2: 17.73
%, eigen alue 1.419; Fig. 5A; Table 3). Plo ing z-sco es
o he indi iduals’ cha ac e s agains he axes de ined
by Fac o s 1 and 2 also e ealed h ee sepa a ed g oups
co esponding o he sampled indi iduals o each species,
Pančo e al.: Pinguicula pan e iae (Len ibula iaceae), a new species om Cuba390
wi h no o e lap in hei mo phospaces (Fig. 5B). Fac o
1 was de ined by ep oduc i e cha ac e s wi h high
posi i e weigh . This componen sepa a ed species based
on in lo escence scape leng h (SCL), lowe leng h (FLL),
and he leng h (LBL, MLL) and wid h (LBW, MLW) o he
co olla lobes. Pinguicula ja aguana appea s o he igh
end o he axis, showing longe in lo escences and la ge
lowe s, ollowed by P. pan e iae, and inally P. albida,
wi h sho e in lo escences and smalle co ollas (Fig. 5B;
Table 2). These ep oduc i e ai s ha e been highligh ed
as ele an cha ac e is ics o disc imina e among ela ed
species o Pinguicula (Con i and Pe uzzi 2006; Pe uzzi
and Ges i 2013; Domínguez e al. 2014; Bu elo-Ramos e
al. 2018; C espo e al. 2020). On he o he hand, Fac o 2
was de ined by lea leng h (LFL) and wid h (LFW), bo h
showing high posi i e weigh (Fig. 5A) indica ing he
p esence o smalle lea es in P. pan e iae; howe e , he
o e all a ia ion o hese ege a i e cha ac e s is a ibu ed
o in aspeci ic a iabili y (Fig. 5B). Mo eo e , quali a i e
ege a i e cha ac e s, such as lea g ow h, shape and
ma gin, we e also ecognised as diagnos ic cha ac e s
use ul o di e en ia e he h ee species (Fig. 4; Table 2),
as p e iously no ed o o he g oups o ela ed axa wi hin
Pinguicula (Caspe e al. 2020; C espo e al. 2020; Pé ez e
al. 2023). These ai s may be associa ed wi h he speci ic
en i onmen al condi ions in which each species h i es,
including hei adap a ion o di e en ligh exposu e and
wa e a ailabili y.
The new species was disco e ed along he banks
o he Ri e Toa wi hin he Alejand o de Humbold
Na ional Pa k. This egion is well-known o i s high
le el o endemism, o en associa ed wi h he p esence
o se pen ine-de i ed soils, and has been highligh ed as
he main cen e o di e si ica ion and endemism o he
moun ainous lo a o Cuba (López Almi all 2013; López
Almi all and Dua e 2022). Pinguicula pan e iae sha es
i s habi a wi h P. ja aguana in e i ic soils de i ed
om se pen ine bed ocks (cha ascales) in eas e n
Cuba; while P. albida occu ences include popula ions
in se pen ine de i ed soils in wes e n Cuba (cuabales)
(Fig. 4). Acco ding o Bo hidi (1996), se pen inophilous
lo as o eas e n and wes e n Cuba esul ed om in ensi e
local specia ion and ep esen independen e olu iona y
scena ios.
Wi h he desc ip ion o P. pan e iae, ou indings
con i m ha Cuba is a ho spo o Pinguicula di e si y,
hos ing a p esen 15 species and one subspecies, all
o which a e endemic o he Cuban a chipelago. This
di e si y aligns wi h he concep o a Cen al Ame ican-
Ca ibbean cen e o di e si y o he genus (Caspe 2019)
and ein o ces he need o a mo e accu a e assessmen
o he axonomic di e si y o he g oup in emo e a eas.
Ten o he Cuban species a e ound exclusi ely in eas e n
Cuba, a egion ecognised as a cen e o endemism o
he genus (Domínguez e al. 2024). Al hough mos Cuban
species a e dis ibu ed wi hin conse a ion uni s, hey
emain exposed o h ea s due o he limi ed geog aphic
anges o known popula ions, pa icula ly hose species
ha a e known om a single loca ion, such as P. pan e iae.
Fu he mo e, as o o he species, he single popula ion o
P. pan e iae may be impac ed by local and global clima e
changes, bu no e idence o speci ic clima e isks o his
axon has been ob ained as ye .
ACKNOWLEDGEMENTS
We a e g ea ly indeb ed and o e ou since e hanks o
se e al indi iduals and o ganisa ions. U li Manuel Bo ges
Calzadilla ob ained he ini ial pe mi s o isi La Melba
and con ibu ed his excep ional assis ance o o ganising
and helping du ing all ield expedi ions. The scien i ic
depu y di ec o o he Unidad de Se icios Ambien ales
Alejand o de Humbold , Ge a do Begué Quiala, g an ed
he necessa y pe mi s and p o ided ield guides. In
pa icula : Jesús Imbe Planche (Head o Oji o de Agua
sec o ) ac ed as guide o loca e he popula ion, while Jo ge
Luis Delgado Labañino (Head o La Melba sec o ) scaled
wa e alls, opened pa hs o almos inaccessible loca ions,
and sha ed his amazing knowledge o local biodi e si y.
Addi ionally, Luis Manuel ‘Güicho’ Velasquez Za agoza
(d i e o La Melba sec o ) and Roge Rosabal Reyes
(d i e o he Unidad de Se icios Ambien ales Alejand o
de Humbold ) demons a ed excep ional d i ing skills as
hey nego ia ed nea ly inaccessible e ain. The he ba ium
o he Ma in-Lu he -Uni e si ä Halle-Wi enbe g
(HAL), in pa icula Denise Ma x, loca ed specimens
and sha ed high-quali y scans and in o ma ion. The
he ba ium o he Cuban Na ional Bo anic Ga den,
Uni e si y o Ha ana (HAJB), in pa icula Eldis R.
Bécque , and he he ba ium o he Cen o O ien al de
Ecosis emas y Biodi e sidad (BSC), in pa icula Ángel
Mo i o Ma ín, Mayelín Silo Ley a, Rosa M. B ooks
La e deza and Yenisey Re illa Góngo a, assis ed in he
apid p ocessing o specimens. Finally, we hank Dasmiliá
C uz and Tilo Henning o hei cons uc i e commen s
and sugges ions on he ea ly e sion o he manusc ip .
REFERENCES
Alain H (1957) Len ibula iaceae. In: León H, Alain H (Eds)
Flo a de Cuba IV. Dico iledóneas: Melas oma aceae a
Plan aginaceae. Con ibuciones Ocasionales del Museo
de His o ia Na u al del Colegio “De La Salle”. Imp eso P.
Fe nández y Cía, La Habana, 472–480.
Bachman S, Moa J, Hill AW, de la To e J, Sco B (2011)
Suppo ing ed lis h ea assessmen s wi h GeoCAT:
geospa ial conse a ion assessmen ool. ZooKeys 150: 117–
126. h ps://doi.o g/10.3897/zookeys.150.2109
Bisse J, Lippold H, Caspe SJ (1975) Bei äge zu Kenn nis
de wes indischen Pinguicula-A en. Wissenscha liche
Zei sch i de F ied ich-Schille -Uni e si ä Jena,
Ma hema isch-Na u wissenscha liche Reihe 24(4): 377–
385.
Bo hidi AL (1996) Phy ogeog aphy and Vege a ion Ecology o
Cuba. Akadémiai Kiadó, Budaspes , 1–923.
Bu elo-Ramos CM, Zamudio-Ruiz S, González-Aguila MA
(2018) Pinguicula olmeca (Len ibula iaceae) una nue a
especie del su de México. Bo anical Sciences 96(2): 359–
365. h ps://doi.o g/10.17129/bo sci.1927
