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Three new species of Monstera (Araceae: Monsteroideae: Monstereae) from the Cordillera de Talamanca in Costa Rica, threatened by the expansion of coffee plantations

Wiley
Nordic Journal of Botany
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Abstract and Figures

Monstera alfaroi Croat & M. Cedeño, M. tarrazuensis Croat & M. Cedeño and M. wilsoniensis M. Cedeño & Grayum, all apparently endemic to central and southern Costa Rica at middle elevations on the Pacific slope, are described as new species. Monstera alfaroi is characterized by having warty light brown petioles with undulate sheaths, very large adult leaves and inflorescences with warty peduncles, and externally light green spathes. Monstera tarrazuensis is characterized by its smooth, light green petioles with white dots and deciduous sheaths, coriaceous, entire leaf blades and inflorescences on hanging stems. Monstera wilsoniensis is characterized by its deciduous petiole sheaths, cordate or subcordate, weakly or not fenestrated leaf blades with entire margins and numerous primary lateral veins, and inflorescences in ascending or hanging stems. The three novelties are compared, respectively, with M. buseyi Croat & Grayum, M. luteynii Madison and M. adansonii Schott, and all are illustrated with color photos of living material.
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NORDIC JOURNAL OF
BOTANY
Nordic Journal of Botany
1
––––––––––––––––––––––––––––––––––––––––
© 2020 Nordic Society Oikos. Published by John Wiley & Sons Ltd
Subject Editor: Bertil Ståhl
Editor-in-Chief: Torbjörn Tyler
Accepted 6 November 2020
2020: e02970
doi: 10.1111/njb.02970
2020 e02970
Published 29 December 2020
Monstera alfaroi Croat & M. Cedeño, M. tarrazuensis Croat & M. Cedeño and M.
wilsoniensis M. Cedeño & Grayum, all apparently endemic to central and southern
Costa Rica at middle elevations on the Pacic slope, are described as new species.
Monstera alfaroi is characterized by having warty light brown petioles with undulate
sheaths, very large adult leaves and inorescences with warty peduncles, and exter-
nally light green spathes. Monstera tarrazuensis is characterized by its smooth, light
green petioles with white dots and deciduous sheaths, coriaceous, entire leaf blades and
inorescences on hanging stems. Monstera wilsoniensis is characterized by its decidu-
ous petiole sheaths, cordate or subcordate, weakly or not fenestrated leaf blades with
entire margins and numerous primary lateral veins, and inorescences in ascending or
hanging stems. e three novelties are compared, respectively, with M. buseyi Croat
& Grayum, M. luteynii Madison and M. adansonii Schott, and all are illustrated with
color photos of living material.
Keywords: Araceae, Central America, conservation
Introduction
Monstera Adans. is a genus of 48 currently-recognized species (but estimated to have
ca 80 species; Boyce and Croat 2011 onwards) of mostly nomadic climbers occurring
in the humid Neotropics at low to medium elevations, and is the only American genus
of the pantropical Rhaphidophora-clade (Zuluagaetal. 2019). It is perhaps best known
for its perforate leaves (which have made many species popular as garden and indoor
plants), though they are not characteristic of all Monstera species or conned to this
genus (see Hay 2019 for a summary of the taxonomic distribution of intra-marginally
dissected (i.e. perforate or fenestrate) leaves in Araceae).
Monstera was taxonomically revised by Madison (1977), who adopted a very broad
species concept, and at least some of his species are now better regarded as complexes.
Grayum (2003b) provided a synopsis for the Costa Rican species, but acknowledged
Three new species of Monstera (Araceae: Monsteroideae:
Monstereae) from the Cordillera de Talamanca in Costa Rica,
threatened by the expansion of coffee plantations
MarcoCedeño-Fonseca, Michael H.Grayum, Thomas B.Croat and Mario A.Blanco
M. Cedeño-Fonseca (https://orcid.org/0000-0003-0119-0427 (marcovf.09@gmail.com), Programa de Posgrado en Biología/Herbario Luis Fournier
Origgi (USJ), Univ. de Costa Rica, San José, Costa Rica. – M. H. Grayum (https://orcid.org/0000-0002-9501-1161) and T. B. Croat (https://orcid.
org/0000-0002-1804-0532), Missouri Botanical Garden, St. Louis, MO, USA. – M. A. Blanco (https://orcid.org/0000-0001-7369-5411), Jardín
Botánico Lankester, Univ. de Costa Rica, Cartago, Costa Rica, and: Centro de Investigación en Biodiversidad y Ecología Tropical, and Escuela de Biología,
Univ. de Costa Rica, San José, Costa Rica.
Research
2
that his treatment was still somewhat tentative. e rst
author has recently completed an as-yet-unpublished revi-
sion for Costa Rica, where Monstera is particularly diverse
and rich in species (Cedeño-Fonseca 2019). In the course
of that revision, some 10 species new to science were iden-
tied (plus several others that were either resurrected from
synonymy or documented as new records for the country).
ree of those new species are described below; two of them
are under imminent threat from the encroachment of coee
plantations. Some other novelties have been published else-
where (Cedeño-Fonsecaetal. 2018, 2020b, c, d, e).
e Cordillera de Talamanca, one of the most oristically
diverse areas in Costa Rica, runs through the center of the
country from the Cerros de Escazú and Cerro Turrubares
southeastward, in a series of peaks exceeding 3000 m a.s.l.,
to the center of Panama, with an average elevation of 1300
m (Hammeletal. 2004, Monroetal. 2017). e Cordillera
de Talamanca in the region of Santa María de Dota and San
Marcos de Tarrazú has experienced a wide expansion of cof-
fee cultivation, reducing the remnants of its natural forests.
ese forests belong to the ‘premontane moist forest’ life
zone, one of the most threatened ecosystems in the country
(Rodríguez and Brenes Cambronero 2009, Cascante-Marín
and Estrada-Chavarría 2012).
‘Premontane moist forests’ are located approximately
between 700 and 1500 m a.s.l. (Bolaños and Watson 1993),
fragmented by urban growth and mainly by coee produc-
tion (Rodríguez and Brenes Cambronero 2009, Cascante-
Marín and Estrada-Chavarría 2012). e original coverage
of the ‘premontane moist forest’ has almost completely dis-
appeared in the region, and it is estimated that only 21%
of the original area currently exists in the country, with a
high degree of fragmentation (Cascante-Marín and Estrada-
Chavarría 2012). However, in these ecosystems an impor-
tant set of endemic species that are not distributed in other
mountainous areas is conserved, e.g. Agave wercklei F.A.C.
Weber ex Wercklé (Grayum 2003a), Calathea tarrazuensis H.
Kenn. (Kennedy 2011), Chamaedorea piscifolia Hodel, G.
Herrera & Cascante (Hodelet al. 1997), Cupania morale-
sii J.E. Jiménez, P. Juárez & J.M. Chaves (Jiménez et al.
2016), Drymonia peltata (Oliv.) H.E. Moore (Kriebel 2010),
Lepanthes dotae Endrés ex Luer (Luer 2003), L. tarrazuensis
Bogarín & D. Jiménez (Bogarínetal. 2012) and Monstera
epipremnoides Engl. (Engler 1905).
During eld explorations in 2018 and 2019 observations
were made of the uncontrolled expansion of coee crops
along the TarrazúQuepos route. ese reach the highest part
of the mountains, without conserving remnants that allow
connectivity between the dierent forest areas, while also
exposing extensive areas of bare soil to erosion and large land-
slides. Monocultures of oil palm and pineapple have spread
exponentially, taking over regions with high biodiversity
(Beggs and Moore 2013, Acuña-Piedra and Quesada-Román
2017, Cedeño-Fonsecaet al. 2020a). However, these have
been established on land with slopes of less than 20°, whereas
coee crops have developed on very steep slopes, of up to
70°. Unfortunately, changes in land use are very poorly regu-
lated by relevant institutions in Costa Rica, with the result
that biodiversity continues to be under signicant threat in
this very rich area.
Monstera has a diversity hotspot in the Cordillera de
Talamanca (Madison 1977). During eldwork in the region
of Santa María de Dota and San Marcos de Tarrazú, two new
endemic species were found in the remnants of forests iso-
lated by the expansion of coee crops. A third new species
that is distributed along the Cordillera de Talamanca has a
population conserved in the Chalchuapa municipal forest,
near Santa María de Dota, though it has not been docu-
mented in forest remnants among coee crops. e popula-
tions of the three new species of Monstera described here in
the area of Santa María de Dota and San Marcos de Tarrazú
are threatened by the expansion of coee crops.
Material and methods
Collections were made in dierent sectors of the Pacic slope
of the Cordillera de Talamanca, and compared with mate-
rial deposited in the herbaria CR, CR-INB, F, HLDG, MO,
NY, PMA, SEL and USJ, and also with images of type mate-
rial of Central American and Colombian species deposited in
other herbaria. e descriptive terminology mostly follows
Madison (1977). e acronyms of all herbaria mentioned are
according to iers (2020), with the exception of CR-INB,
which denotes specimens deposited in the former INB her-
barium, now nominally incorporated in CR but still housed
in the o-site building, formerly of the Instituto Nacional de
Biodiversidad, in Santo Domingo de Heredia, Costa Rica.
In the descriptions below, the term ‘nomadic vine’ (or
‘nomadic appressed climber’) follows Sperottoetal. (2020).
e seedling stage is dened as that after germination but
before the plant starts to climb a phorophyte (host tree); the
juvenile stage begins at that point, and the adult stage begins
when the foliage leaves attain the same general shape and
size as that of reproductive plants. e juveniles of the three
new species treated here (which are members of Monstera sec-
tion Monstera, sensu Madison 1977) are described as bearing
foliose leaves, to emphasize the dierence with the stolon-
iferous seedlings that bear scale leaves, of species in section
Echinospadix Madison and section Marcgraviopsis Madison
(Madison 1977).
Each of the three species described below is illustrated by
a photographic plate of dierent parts of the plants in situ,
adapted from the ‘Lankester Composite Dissection Plate’
technique developed for Orchidaceae at Lankester Botanical
Garden, Univ. de Costa Rica (Pupulin and Bogarín 2004).
Results
Monstera alfaroi Croat & M. Cedeño, sp. nov. (Fig. 1, 2)
A species characterized by having warty, light brown petioles,
deciduous petiole sheaths, scarcely fenestrated and subcoria-
ceous leaf blades, and inorescences with warty peduncles
and the spathe light green externally.
3
Type: Costa Rica, San José: Tarrazú county, San Lorenzo
district, road from Tarrazú to Quepos, 9°3457,47N,
84°42,53W, 1400 m, 22 Oct 2019, M. Cedeño 1702
(holotype USJ!; isotype MO!, to be distributed).
Etymology
e name alfaroi is given in honor of the Costa Rican bota-
nist Evelio Gerardo Alfaro Vindas from the Las Mellizas area
of Puntarenas province, Costa Rica, who made one of the
earliest herbarium collections of this species.
Description
Robust nomadic appressed-climber. Seedlings bearing foliose
leaves. Juvenile plants appressed-climbing. eir stem dark
green, cylindrical or dorsally attened, with whitish-green
pustules making a rough-warty surface; internodes 1–15 cm
Figure1. Monstera alfaroi sp. nov. (A) developing inorescence, (B) portion of juvenile plant, (C) inorescence at female anthesis; spathe
cut longitudinally to show spadix inside, (D) stylar cap with stigma (left) and one stamen (right), (E) fertile ower, in lateral view (left), and
longitudinal section (right), (F) apical portion of adult plant, (G) sterile ower, in lateral view (left), and longitudinal section (right). Based
on M. Cedeño 1702 (USJ) in life.
4
long × 5–10 mm diam. Petioles partly concealed by the leaf
blades or not at all, light green, warty, 3–16 cm long, sheathed
to the base of the geniculum; petiole sheath deciduous; leaf
blades not appressed to the phorophyte, ovate-lanceolate,
acuminate at apex, coriaceous, 9–20 × 8–13 cm, without
fenestrations.
Adult plants appressed-climbing. eir stem dark green,
warty, cylindrical; internodes 1–2 cm long × 2–3 cm diam.
Support and feeder roots unknown. Petioles light green and
light brown at the base, with black warts, 40–70 cm long,
sheathed along their entire length; petiole sheath undulate,
persistent or deciduous; geniculum warty throughout, at-
tened adaxially and transversally convex abaxially, 2.0–2.5
cm long. Leaf blades narrowly ovate, rounded or asymmetric
at base, shortly acuminate at apex, subcoriaceous, with entire
margins and few perforations, 60–90 × 30–45 cm, drying
dark brown and weakly lustrous, not decurrent on the genicu-
lum; midrib concave adaxially, convex abaxially, drying black
or yellowish; primary lateral veins 8–13 per side, diverging
at 50–70°, impressed adaxially, prominent abaxially; tertiary
Figure2. Monstera alfaroi sp. nov. (A) adult plant in its natural habitat, (B) leaves of adult plant (up to 190 cm long) with Mick Mittermeier
and Marco Cedeño for scale, (C) detail of petiole base, showing black warts and light brown epidermis (arrow), (D) cataphyll (left arrow)
and prophyll (right arrow), both with black and white pustules and undulate margins.
5
veins parallel to the primary lateral veins; collecting veins
barely visible. Inorescences produced on ascending stems;
peduncle warty, 20–35 cm long; spathe obtuse or acuminate,
light green in developing inorescences, internally white and
externally light green at anthesis, 13–20 × 8–14 cm; spa-
dix white during development, white or yellowish white at
anthesis, 10.0–15.0 × 1.2–2.5 cm. Sterile owers 4–5 mm
long; fertile owers 5–7 mm long. Stamens with laminar
laments 2–7 mm long; anthers 2.0–2.5 mm long. Ovary
squarish, 2.0–3.0 × 1.5–2.5 mm; style hexagonal, 1.2–2.0 ×
2.0–3.0 mm long; stigmatophore slightly columnar; stigma
linear with transparent secretion. Fruits with the stylar cap
after anthesis white, becoming white-cream when ripe; pulp
white; seeds black, 3–6 mm long.
Distribution and ecology
Endemic to Costa Rica in the central and south Pacic
regions, in the provinces of Puntarenas and San José, from
Tarrazú south to Potrero Grande and the Fila Costeña, at
1100–1500 m a.s.l. (Fig. 7). Inhabits primary and secondary
forests, on trees usually with diameters greater than 45 cm.
Phenology
Flowering has been recorded in February, October and
November and fruiting in January.
Conservation status
Monstera alfaroi is safely conserved in a small geographic area
of the Parque Internacional de La Amistad. e habitat where
it grows is strongly impacted by human activities, especially
due to agricultural expansion. We consider that it qualies
for the endangered EN [B1ab(ii, iii, iv) + B2ab(ii, iii, iv)] cat-
egory, according to the IUCN Red List Criteria.
Note
Monstera alfaroi diers from other species of its genus in
Costa Rica by having light brown petioles with black warts
throughout their length, undulate and warty petiolar sheaths
that become deciduous, coriaceous and scarcely fenestrated
leaf blades, warty inorescence peduncles and spathes light
green externally. Adult leaves of M. alfaroi are among the
largest of Costa Rican species of Monstera section Monstera
(Fig. 2B); as with other species in the genus, the leaves adja-
cent to the inorescences are smaller than others on the same
plant (Fig. 1). We have not yet been able to document a prop-
erly open spathe of M. alfaroi in life, so its description above
is incomplete and based on the notes in the available ower-
ing herbarium specimens.
Monstera alfaroi is most closely related to M. buseyi Croat
& Grayum, with which it shares tuberculate petioles. Most
collections of M. buseyi dier from M. alfaroi in having nar-
rowly ovate-elliptic leaf blades 2.0–2.5 times longer than
wide which dry medium grayish yellow-brown adaxially and
much paler (yellowish green) and often matte abaxially (ver-
sus leaf blades dry dark brown but with blades only 1.7 times
longer than wide in M. buseyi), although some collections
of M. alfaroi have blades only 1.7 times longer than wide
that dry dark brown. Also, M. buseyi has owering spadices
only 6–9 cm long and ca 1 cm in diameter, versus more than
18 cm long and 2 cm in diameter in M. alfaroi. Finally, M.
buseyi has inorescence peduncles drying matte and smooth
or only weakly puberulent-scurfy, whereas those of M. alfa-
roi dry grossly ridged and pitted and covered by conspicuous
tubercles with the surfaces semiglossy.
Monstera alfaroi can also be confused with M. costaricen-
sis (Engl. & K. Krause) Croat & Grayum, but the latter has
warty stems, petioles with white pustules and markedly undu-
late persistent sheaths, and owers with markedly conical
styles. Monstera alfaroi is restricted to the Pacic slope of the
Cordillera de Talamanca, while M. costaricensis occurs only in
the Caribbean lowlands (up to 1050 m a.s.l.) of Nicaragua,
Costa Rica and Panama (Cedeño-Fonsecaetal. 2020b).
Monstera alfaroi was treated as ‘Monstera sp. M’ in Cedeño-
Fonseca (2019).
Additional specimens examined (paratypes)
Costa Rica, Puntarenas: Buenos Aires, Potrero Grande, La
Lucha, Punto 8, 9°626.8N, 83°546.56W, 1450 m, 20
Feb 2018, D. Santamaríaetal. 7036 (MO); Parrita, Parrita,
Fila Chonta, 9°3527N, 84°3630W, 1300 m, 27 Nov
2003, A. Quesada 1264 (CR); Parrita, Fila Chonta, road
from San Marcos de Tarrazú to Cerro Cura, La Virgen
and Fila Chonta, 9°350N, 84°100W, 1400 m, 31 Jan
1996, B. Hammel & J. Morales 21189 (CR-INB, MO).
San José: Tarrazú, San Lorenzo, Nápoles, ridges to the west
of Cerro Toro, 9°3430N, 84°435W, 1200 m, 30 Nov
1995, G. Herrera et al. 8785 (CR); Pérez Zeledón, Fila
Costeña, 9°1741N, 83°4551W, 1100 m, 31 Jan 1998,
B. Hammel et al. 20161 (CR-INB, MO); Pérez Zeledón
county, Cordillera de Talamanca, Santa Elena station, 1250
m, 15 Feb 1996, E. Alfaro 508 (CR, MO).
Monstera tarrazuensis Croat & M. Cedeño, sp. nov.
(Fig. 3, 4)
A species characterized by its light green petioles with white
dots, sheathed along their entire length, with deciduous peti-
ole sheaths, coriaceous leaf blades entire and without fenes-
trations, inorescences produced on hanging stems, spathe at
anthesis whitish-green externally and white internally, with
margins overlapping toward the base, and fruits creamy when
ripe and with white pulp.
Type: Costa Rica, San José: Tarrazú county, San Lorenzo
district, road from Tarrazú to Quepos, 9°3457,47N,
84°42,53W, 1400 m, 9 Jul 2019, M. Cedeño 1686 (holo-
type USJ!).
Eponymy
e epithet tarrazuensis refers to the Tarrazú Cantón (county)
in San José Province, where the type locality is found.
6
Description
Nomadic appressed-climber. Seedlings bearing foliose leaves.
Juvenile plants appressed-climbing. eir stem light green,
mottled with white smooth dots; internodes 3–7 cm long ×
3–5 mm diam.; petioles not concealed by the leaf blades, dark
or light green, smooth, 4–10 cm long, sheathed to the base of
the geniculum or to the base of the leaf blade; petiole sheath
persistent; leaf blades not appressed to the phorophyte, lan-
ceolate, cordate at the base, acuminate at apex, coriaceous,
7–14 × 5–9 cm, without fenestrations.
Adult plants appressed-climbing. ier stem dark green
with white dots, smooth, cylindrical; internodes 4.0–11.0 cm
long × 1.2–1.7 cm in diam. Support and feeder roots light
brown. Petioles light green with white dots, smooth, 11–14
cm long, sheathed along their entire length; petiole sheath
deciduous; geniculum 0.5–1.0 cm long; leaf blades ovate to
ovate-elliptic, asymmetrically rounded or subtruncate at base,
shortly acuminate at apex, coriaceous, 13.0–26.0 × 7.5–13.8
cm, not decurrent on the geniculum, drying gray or brown,
glossy, without perforations, with entire margins; midrib
Figure3. Monstera tarrazuensis sp. nov. (A) juvenile plant, (B) adult hanging stem, (C) sterile ower, in lateral view (left), and longitudinal
section (right), (D) fertile ower, in lateral view (left), and longitudinal section (right), (E) developing infructescence, (F) stylar cap with
stigma (left) and one stamen (right), (G) developing inorescence. Based on M. Cedeño 1686 (USJ) in life.
7
concave adaxially, convex abaxially, drying dark brown-black;
primary lateral veins 5–9 per side, diverging at 45–60°, sig-
moid, impressed adaxially, prominent abaxially, drying dark
brown; tertiary veins parallel to the primary lateral veins;
collective veins barely visible. Inorescences produced on
hanging stems; peduncle smooth, 13–15 cm long; spathe
long-acuminate, the margins overlapping at the lower half,
light green in developing inorescences, green whitish exter-
nally and white internally at anthesis, 10–15 × 5–7 cm; spa-
dix white, 5–6 × 2–3 cm. Sterile owers 2.5–4.0 mm long,
with transparent secretion; fertile owers 3–4 mm long.
Stamens with laminar lament 2–4 mm long; anthers 1.5–
2.0 mm long. Ovary rectangular in longitudinal section, 2.0–
3.0 × 1.5–2.0 mm; style hexagonal, 1.5–2.0 × 2.0–3.0 mm,
without stigmatophore; stigma linear, with transparent secre-
tion. Fruits with the stylar cap after anthesis white, becoming
white-cream when ripe; pulp white; seeds unknown.
Distribution and ecology
Endemic to Costa Rica in the central Pacic region, on and
near Cerro Hormiguero, at 1100–1386 m a.s.l. (Fig. 7).
Inhabits primary and secondary forests.
Phenology
Flowering has been recorded in July, and fruiting July and
September.
Conservation status
Monstera tarrazuensis is not found in any protected areas.
e habitat where it grows is strongly impacted by human
activities, especially the expansion of coee plantations. We
consider it qualies as critically endangered CR [B1ab(ii,
iii, iv) + B2ab(ii, iii, iv)], according to the IUCN Red List
Criteria.
Note
Monstera tarrazuensis diers from other species of the genus
in Costa Rica by its smooth, pale green petioles with white
dots, petiole sheaths deciduous, small (13.0–26.0 × 7.5–13.8
cm), coriaceous leaf blades, and inorescences produced only
on hanging stems, with the margins of the spathe overlap-
ping in the lower half. Monstera tarrazuensis can be confused
with M. luteynii Madison, which also has unperforated leaf
blades and produces inorescences on hanging stems, but it
diers because M. luteynii has stems covered with pustules,
Figure4. Monstera tarrazuensis sp. nov. (A, B) adult plants in their natural habitat, with hanging stems at the edge of the forest, (C) hanging
stem with erect inorescence (arrow), (D) hanging stem with erect infructescence (arrow).
8
and leaves with terete genicula and blades with tertiary veins
strongly prominent on both surfaces.
Monstera tarrazuensis and M. luteynii are both species
with a hanging habit (i.e. with inorescences produced on
hanging stems) at elevations above 1100 m a.s.l. Other Costa
Rican species of Monstera with hanging habit are M. molinae
Grayum, M. pittieri Engl. and M. tuberculata Lundell.
Monstera tarrazuensis diers from M. molinae in having
leaf blades with entire margins and without fenestrations
(versus leaf blades pinnatilobed), fruits with the stylar cap
white-cream when ripe (versus moss-green). It diers from
M. pittieri in having long-acuminate spathes (versus obtuse
or mucronate), fragmented deciduous petiolar sheaths (ver-
sus partly persistent with brous residues), and fruits with the
stylar cap white-cream when ripe (versus yellow).
Monstera tarrazuensis was treated as ‘Monstera sp. N’ in
Cedeño-Fonseca (2019).
Additional specimens examined (paratypes)
Costa Rica, San José: Tarrazú, San Marcos de Tarrazú
between Cerro Toro and Cerro Hormiguero along the road
between Basuero de Tarrazú and Esquipulas, 9°3330N,
84°0315W, 1150 m, 5 Sep 1996, T. Croat 78934 (CR-INB,
MO); Tarrazú county, San Marcos de Tarrazú between Cerro
Toro and Cerro Hormiguero along the road between Basuero
de Tarrazú and Esquipula, vicinity of Cerro Hormiguero,
9°3330N, 84°0315W, 1150 m, 5 Sep 1996, T. Croat
78912 (CR, MO).
Monstera wilsoniensis M. Cedeño & Grayum,
sp. nov. (Fig. 5, 6)
A species characterized by its petioles sheathed throughout
their length, with the petiole sheath deciduous, leaf blades
cordate or subcordate to obtuse at the base, weakly fenestrate,
with entire margins and numerous primary lateral veins,
inorescences borne on both ascending and hanging stems,
with the spathe becoming completely open, long-acuminate
or mucronate, externally yellowish green and internally white,
and owers without stigmatophores and with linear stigmas.
Type: Costa Rica, Puntarenas: Coto Brus, San Vito, Jardín
Botánico Wilson, forest preserve, 8°471N, 82°5858W,
1350 m, 10 Oct 2018, M. Cedeño, M. Blanco, M. Mata &
O. Alvarado 1484 (holotype, USJ [2 sheets]; isotypes, MO,
PMA, to be distributed).
Eponymy
e epithet wilsoniensis refers to the Wilson Botanical Garden
(part of the Reserva Biológica Las Cruces, owned by the
Organization for Tropical Studies; Bates 1987), a site where
this species is especially abundant.
Description
Nomadic appressed-climber. Seedlings bearing foliose leaves.
Juvenile plants appressed-climbing. eir stem light green,
with black pustules; internodes 3–5 cm long × 4–6 mm diam.
Petioles not concealed by leaf blades, dark or light green,
smooth, 5–10 cm long, sheathed to the base of the genicu-
lum; petiole sheath deciduous; leaf blades not appressed to
the substrate of the phorophyte, ovate to lanceolate, attenu-
ate or truncate and asymmetric at base, acuminate at apex,
thinly coriaceous, 8–15 × 2–6 cm, without fenestrations.
Adult plants appressed-climbing. eir stem light green to
dark green, cylindrical, with brown or black pustules, inter-
nodes 2.0–10.0 cm long × 1.0–1.5 cm diam. Support roots
dark brown; feeder roots brown. Petioles dark green, with or
without white dots, smooth, 13–20 cm long, sheathed to the
base of the geniculum; petiole sheath deciduous; geniculum
smooth, adaxially grooved, round abaxially, 1.0–2.5 cm long;
leaf blades ovate to lanceolate, oblique, cordate to obtuse at
base, acuminate at apex, thinly coriaceous, drying greenish,
yellowish or blackish, 15–25 × 13–15 cm, slightly decurrent
to the middle of the geniculum, with entire margins, fenes-
trations few or none; midrib concave adaxially, convex abaxi-
ally, drying yellowish or blackish on both surfaces; primary
lateral veins 15–25 per side, bifurcated, diverging at 55–65°,
impressed adaxially, prominent abaxially, drying yellowish or
blackish; tertiary veins reticulated; collecting veins evident.
Inorescences produced on both ascending and hanging
stems; peduncle smooth, 10–15 cm long; spathe long-acumi-
nate or mucronate, light green in developing inorescences,
at anthesis externally yellowish green, internally white, rela-
tively thin, open throughout, deciduous after anthesis, 13–18
× 9–13 cm; spadix in development cream-colored, at anthesis
yellowish cream, 7.0–13.0 × 1.5–2.0 cm. Sterile owers 3–5
mm long, with transparent secretion; fertile owers 4–6 mm
long. Stamens with laminar laments 2–6 mm long; anthers
1–2 mm long. Ovary square, 3–4 × 3–4 mm; style hexago-
nal, 1–2 × 3–4 mm, without stigmatophore; stigma linear,
with transparent secretion. Fruits with the stylar cap green
after anthesis, becoming white-cream when ripe; pulp white;
seeds black, 3–5 mm long.
Distribution and ecology
Monstera wilsoniensis is seemingly endemic to Costa Rica on
the Pacic slope of the Cordillera of Talamanca, as well as the
Cerros de La Carpintera, Cerros de Escazú and Fila Costeña,
from 900 to 2200 m a.s.l., in ‘premontane moist forest’ life
zone (Fig. 7). Because it has been collected a few km from the
Panamanian border, it is expected to eventually turn up in
adjacent Chiriquí province in Panama.
Phenology
Flowering has been recorded from March to October, fruit-
ing throughout the year.
Conservation status
Monstera wilsoniensis is protected at the Estación Biológica
Las Cruces, on the Pacic versant of La Amistad International
Park, and in the Cerros de La Carpintera and Cerros de
Escazú Protected Areas. However, these protected areas
are strongly impacted by human activities, especially due
to agricultural expansion. We considered it qualies as an
9
endangered species, EN [B1ab(ii, iii, iv) + B2ab(ii, iii, iv)]
category according to the IUCN Red List Criteria.
Note
Monstera wilsoniensis diers from other species of the genus
in Costa Rica by having petioles sheathed along their entire
length, petiole sheaths deciduous, leaf blades usually cor-
date, scarcely or not fenestrate, with entire margins and
with numerous lateral primary veins, and inorescences
produced on both ascending and hanging stems. is spe-
cies has been confused with M. adansonii Schott (specically
with the morphotype that occurs on the Pacic lowlands of
Costa Rica), but diers from the latter by having shorter
petioles (13–20 versus 35–55 cm) and smaller leaf blades
(15–25 × 13–15 cm versus 25–65 × 25–45 cm) that are
cordate or subcordate to obtuse (versus cuneate or attenu-
ate) at the base. Monstera wilsoniensis grows mainly in the
‘premontane moist forests’ life zone of the Cordillera Central
Figure5. Monstera wilsoniensis sp. nov. (A) developing inorescence, (B) frontal (left) and back (right) views of open inorescence, (C)
infructescence, (D) fertile ower, in lateral view (left), and longitudinal section (right), (E) stylar cap with stigma (left) and one stamen
(right), (F) portion of juvenile plant, (G) portion of adult plant, (H) sterile ower, in lateral view (left), and longitudinal section (right).
Based on M. Cedeñoetal. 1484 (USJ) in life.
10
and Cordillera de Talamanca, at elevations of 900–2200 m,
while the M. adansonii morphotype that could be confused
with it grows in ‘tropical dry forest’ and ‘tropical moist for-
est’ life zones at elevations of 0–900 m and has conspicu-
ously fenestrate leaf blades.
One of the paratypes, Dodge & Goerger 9825 (MO)
appears to have mixed-up locality data; the given coordinates
and elevation coincide with the province and county (San
José, Dota, 1146 m), but the rest (‘manglar on estero between
Río Paquita and Río Viejo’; compare with ‘Paquita, Río’ in the
Gazetteer of Costa Rican Plant Collecting Locales, <www.
mobot.org/MOBOT/CostaRicaGazetteer/gazetteer.aspx>)
refers to mangrove vegetation on an estuary (obviously at or
near sea level). Given that all other (numerous) collections
have been made at or above 900 m, it is evident that the sec-
ond part of the locality data is erroneous. Coincidentally, this
specimen (collected in 1936) represents the earliest known
herbarium record of this species.
Monstera wilsoniensis was treated as ‘Monstera sp. E’ in
Cedeño-Fonseca (2019).
Additional specimens examined (paratypes)
Costa Rica, Cartago: La Unión, San Diego, La Carpintera
Protected Area, 9°537.27N, 83°5858.72W, 1800 m, 28
Sep 2006, J. Sánchez 1586 (CR); La Unión, San Rafael, La
Carpintera Protected Area, border of the Campo Escuela
Iztarú, 9°533.2N, 83°5846.4W, 1850 m, 13 Jun 2007,
A. Cascante et al. 1754 (CR); La Unión, San Diego, La
Carpintera Protected Area, northern slope, 9°5327.4N,
83°598.9W, 1500 m, 26 Mar 2008, A. Cascanteetal. 1905
(CR); Cartago, San Nicolás, La Carpintera Protected Area,
9°5246N, 83°5813W, 1700 m, 21 Mar 2006, A. Quesada
1604 (CR); La Unión, San Diego, Cerro La Carpintera, slope
toward Tres Ríos, 9°539.5N, 83°5825.8W, 1700 m, 28
Feb 2013, A. Cascante & C. Trejos 2387 (USJ); Cartago.
San Nicolás, Cerros La Carpintera, 9°537N, 83°5748W,
Figure6. Monstera wilsoniensis sp. nov. (A, B) adult plants in their natural habitat, (C) hanging stems with pendulous or sub-erect
infructescences.
11
1700 m, 22 Apr 2016, A. Cascante & C. Trejos 2593 (USJ);
Cartago, San Nicolás, Cerros de La Carpintera, 9°5320N,
83°5810W, 1750 m, 3 Sep 2013, A. Cascante & C. Trejos
2409 (USJ). Puntarenas: Buenos Aires, Biolley, La Amistad
International Park, Cordillera de Talamanca, area around
Río Canasta, 9.5 airline km NW of Agua Caliente, between
Cerro Frantzius and Cerro Pittier, 9°20N, 82°590W,
1550 m, 6 Sep 1984, G. Davidseetal. 28424 (CR, MO);
Coto Brus, San Vito, Las Cruces Biological Station, Java,
forest preserve, 1150 m, 4 Jul 1994, W. Kress & S. Martén
94-4360 (CR); Las Cruces, 8°4724N, 82°570W, 1300
m, 11 Jan 1978, T. Croat 44388 (MO); Coto Brus. San Vito,
Las Cruces Biological Station, Las Cruces Tropical Botanical
Garden, 6 km W of San Vito de Java, 8°490N, 82°580W,
1200 m, 6 Mar 1984, T. Croat 57235 (CR, MO); Coto
Brus, San Vito, Las Cruces Biological Station, along trail
between Las Cruces Botanical Garden and Río Jaba, ca 3.5
km SE of San Vito de Coto Brus, 8°470N, 82°580W,
1200 m, 12 Sep 1985, M. Grayumetal. 5974 (CR, MO);
Coto Brus, San Vito, Las Cruces Biological Station, Wilson
Botanical Garden, Las Cruces, San Vito de Coto Brus, along
eastern trail to Río Java, 8°4730N, 82°580W, 1100 m,
3 Sep 1993, G. Rivera 2151 (CR); Coto Brus, San Vito, Las
Cruces Biological Station, forest below Las Cruces Biological
Station along trail to Rìo Java, San Vito de Java, 8°478N,
82°5729W, 1250 m, 29 Jun 1994, W. Kress & W. Alverson
94-3773 (CR); Coto Brus, San Vito, Las Cruces Biological
Station, Java, forest preserve, 8°478N, 82°5729W,
1250 m, 8 Apr 1994, W. Kress & J. Runk 94-4654 (USJ);
Coto Brus, Pittier, western border of station, 9°142.17N,
82°5753.87W, 1700 m, 2 Aug 2000, L. Acosta 2425 (CR);
Coto Brus, San Vito, Wilson Botanic Garden, Las Cruces,
along eastern trail to Río Java, 8°4730N, 82°580W,
1100 m, 30 Sep 1993, G. Rivera 2163 (CR); Coto Brus,
Las Cruces Biological Station, Ridge Trail, 8°4708N,
82°5811W, 1250 m, 21 Jun 2008, F. Oviedo-Brenes 007
(HLDG); Coto Brus, San Vito, Las Cruces Biological Station,
Loop Trail, 8°4657N, 82°5805W, 1100 m, 17 Sep 2008,
F. Oviedo-Brenes 286 (HLDG); Coto Brus, San Vito, Las
Cruces Biological Station, Rio Jaba Trail at the junction with
Melissa Trail, 8°4713N, 82°5791W, 1150 m, 9 Jul 2009,
F. Oviedo-Brenes 612 (HLDG); Coto Brus, Las Cruces
Biological Station and Wilson Botanic Garden, 6.4 km S of
San Vito on a ridge of the Fila Zapote, Loop Trail, 8°7849N,
82°9676W, 3732 ft, 18 Jan 2007, D.T. Busemeyeretal.
2703 (HLDG); Coto Brus, Las Cruces Biological Station and
Wilson Botanic Garden, 6.4 km S of San Vito on a ridge
of the Fila Zapote, Rio Java Trail, 8°7843N, 82°9611W,
3835 ft, 15 Jan 2007, L.R. Phillippeetal. 39322 (HLDG).
San José: Acosta, Palmichal, Cerros de Escazú Protected Area,
Figure7. Map of Costa Rica showing distributions of Monstera alfaroi (red circles), M. tarrazuensis (green triangles) and M. wilsoniensis
(white triangles).
12
upper watershed of Río Negro, 9°5129.8N, 84°108.3W,
1800 m, 8 Jul 2010, A. Cascanteetal. 2229 (CR); Alajuelita,
San Antonio, Cerros de Escazú Protected Area, around
Cerro Rabo de Mico, 9°5055N, 84°815W, 2200 m, 22
Sep 1989, G. Vargas & J. Sanchéz 765 (CR); Aserrí, Vuelta
de Jorco, Tarbaca, Bajos de Praga, junction of Tarbaca and
Cedral rivers, 9°4810N, 84°730W, 1450 m, 29 Jan
1994, J. Morales 2299 (CR); Desamparados, San Miguel, 4.5
km E of Tobosi, along road 228, 9°5016.6N, 84°135W,
1850 m, 4 Aug 2015, A. Cascante & C. Trejos 2546 (USJ);
Dota, Copey, Providencia, 9°3152N, 83°4948W, 1850
m, 11 Oct 2002, J. Sánchez 1344 (CR); Acosta, Sabanillas,
Bajo Vanegas, around Quebrada Delicias, 9°4152N,
84°1250W, 1500 m, 17 Dec 1996, B. Hammel 20604
(CR); Pérez Zeledón, Páramo, Providencia, Pierdas crossing,
4.7 SE of Providencia, 9°3152N, 83°4948W, 1850 m,
21 Mar 2001, A. Rodríguez 7276 (CR, MO); Dota, between
Río Paquita and Río Viejo, 9°3524N, 83°5548W, 1150
m, 14 Aug 1936, C. Dodge & V.F. Goerger 9825 (MO; with
mixed-up locality data; see note); Dota, Copey, Providencia,
Zapotal, 9°3214N, 83°5018W, 1800 m, 20 Aug 2003,
A. Rodríguez 8262 (CR, MO); Mora, Tabarcia, Cerros
de Escazú Protected Area, watershed of Rio Negro, 2 km
NE of Palmichal, 9°5044.2N, 84°115.8W, 1300 m,
19 Aug 2010, J. Sánchez & R. Chacón 2166 (CR); Pérez
Zeledón, San Pedro, Las Nubes, Santa Elena, 9°2330N,
83°3550W, 1200 m, 4 Aug 1995, E. Alfaro 324 (CR);
Pérez Zeledón, El General, University of York Preserve,
9°2317N, 83°3555W, 1200 m, 28 Nov 2017, M.
Cedeño & M. Mejía 1138 (USJ); Pérez Zeledón, Páramo,
Providencia, Zapotal, 9°3150N, 83°4949W, 1800 m, 20
Aug 2003, A. Ruíz & S. Lobo 743 (CR).
Acknowledgements – MC-F thanks the Missouri Botanical Garden
for an Alwyn H. Gentry Fellowship, the Society of Systematic
Biologists (SSB) for a Mini-ARTS research grant, the Organization
for Tropical Studies for a Glaxo-Wellcome research grant, and the
Rexford Daubenmire fellowship, all in support for the project
‘Taxonomy of the genus Monstera (Alismatales: Araceae), for Costa
Rica’. We thank Mick Mittermeier, Cristina Goettsch Mittermeier
and Caroline Sparks for their help in obtaining funds through a
Gofundme.com campaign. Director Adam Karremans and the sta
at Lankester Botanical Garden (Univ. de Costa Rica) are thanked
for allowing the cultivation of living plants for research. José
Esteban Jiménez and Jairo Jiménez are thanked for their help in
the eld. Alistair Hay is thanked for his help in editing the texts.
We are thankful to the Ministerio del Ambiente y Energía de Costa
Rica (MINAE) and its Sistema Nacional de Áreas de Conservación
(SINAC) for issuing the scientic permits under which wild
specimens were collected. is contribution represents part of the
MSc thesis of Marco Cedeño-Fonseca, completed in the Programa
de Posgrado en Biología at Univ. de Costa Rica.
Author contributions – Marco Cedeño-Fonseca, Michael H. Grayum
and omas B. Croat made plant identications. Marco Cedeño-
Fonseca reviewed herbarium material, took the photographs, and
secured most of the funding. Marco Cedeño-Fonseca, Michael
H. Grayum, omas B. Croat and Mario A. Blanco collected
the specimens. Marco Cedeño-Fonseca and Mario A. Blanco
conceptualized the rst author’s thesis project and wrote the text.
All authors contributed to the design and implementation of the
research and reviewed all versions of the manuscript.
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