Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 649-664, April-June 2021 (Published Jun. 09, 2021)
Phenology of the endangered palm Ceroxylon quindiuense
(Arecaceae) along an altitudinal gradient in Colombia
Blanca Martínez
*; https://orcid.org/0000-0002-7074-3534
René López Camacho
; https://orcid.org/0000-0003-2026-0371
Luis Santiago Castillo
; https://orcid.org/0000-0003-2193-7516
Rodrigo Bernal
; https://orcid.org/0000-0002-9832-8498
1. Universidad Distrital Francisco José de Caldas, Bogotá, Colombia; bamartinezh@correo.udistrital.edu.co
(*Correspondence), rlopezc@udistrital.edu.co
2. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia;
3. Reserva Natural Guadualito, Montenegro, Quindío, Colombia; rgbernalg@gmail.com
Received 27-XI-2020. Corrected 20-III-2021. Accepted 18-V-2021.
Introduction: Understanding the phenology of plant populations is vital for their conservation and management.
We studied the vegetative and reproductive phenology of the endangered palm Ceroxylon quindiuense along an
altitudinal gradient in the Central Cordillera of Colombia. Objective: We describe the leaf production rate, and
flowering and fruiting cycles, and calculate food offer for the fauna, as a tool for the proper management of the
palm. Methods: At each sampling site (2 400, 2 600, 2 800, 3 000 m.a.s.l.), we marked 40 adult individuals (20
pistillate, 20 staminate), which we followed bimonthly for 24 months. We studied leaf production by counting
fallen leaves. We followed flower and fruit production through observations with binoculars and photographs.
Results: Each adult individual produced, on average, one leaf every 69 days. Although isolated individuals
flowered throughout the year, most palms flowered synchronously at each elevation in October 2016-August
2017 and in August 2018-February 2019 and had ripe fruits 7-13 months later. Flowering started at 2 600 m, fol-
lowed by 2 800 and 3 000 m. Palms at 2 400 m, the lower limit of the palm stands in the area, showed a singular
behavior, with scarce flower and fruit production, some individuals that changed sex, and a higher proportion
of pistillate palms. Each palm produced 1-11 (x
= 5.3, SD = 2.2) inflorescences and 1-10 (x
= 5.3, SD = 2.2)
infructescences. The average number of fruits per infructescence was 4 465 (SD = 1 488). With an estimated
population of adult palms between 256 000 and 600 000 and an overall ratio of pistillate: staminate individuals
1:1 or 1:2, total fruit production in the area during each fruiting period is estimated as 2.0-7.1 billion fruits.
Conclusions: The huge number of flowers and fruits and their gradual availability along the altitudinal gradient
have a major impact on the spatial and temporal distribution of food offer for fauna associated with the palm.
Key words: altitudinal gradient; flowering; fruiting; leaf production; palm phenology.
Martínez, B., López Camacho, R., Castillo, L.S., & Bernal,
R. (2021). Phenology of the endangered palm Ceroxylon
quindiuense (Arecaceae) along an altitudinal gradient in
Colombia. Revista de Biología Tropical, 69(2), 649-664.
Palms are an important element in the
dynamics of tropical ecosystems: e.g., by influ-
encing the distribution and behavior of animals,
which could affect other ecosystem functions,
such as plant regeneration (Salm, Jalles-Filho,
& Schuck-Paim, 2005; Beck, 2007; Keuroghl-
ian & Eaton, 2009) and soil properties (Young,
Raab, McCauley, Briggs, & Dirso, 2010).
Palms are a vital food source for many animal
species (e.g., Zona & Henderson, 1989; Peres,
Revista de Biología Tropical, ISSN: 2215-2075 Vol. 69(2): 649-664, April-June 2021 (Published Jun. 09, 2021)
1994; Galetti, Ziparro, & Morellato, 1999;
Scariot, 1999; Genini, Galetti, & Morellato,
2009; Giombini, Bravo, & Tosto, 2016). They
also play an important role for human popu-
lations, which use them as a source of food,
raw construction materials, and for making
a variety of implements, among many other
uses (e.g., Bernal, 1992; Henderson, Galeano,
& Bernal, 1995; Johnson, 2011; Bernal &
Galeano, 2013).
The Quindío wax palm, Ceroxylon quin-
diuense, is an abundant species in the cloud
forests of Colombia and Perú. In Colombia, it
grows in the three Andean ranges, although it
is more abundant in the Central Cordillera. Its
largest populations are located in the Tochecito
River basin, in the municipalities of Cajamarca
and Ibagué, Department of Tolima (Bernal,
Galeano, & Sanín, 2015), where the number
of adults has been estimated between 256 000
(Jiménez, 2017) and 600 000 individuals (Ber-
nal et al., 2015; Instituto Humboldt, 2016).
C. quindiuense is recognized worldwide
for being the world’s tallest palm, reaching
60 m in height (Bernal, Martínez, & Sanín,
2018), and Colombia’s national tree. It is con-
sidered Endangered (EN) (Galeano & Bernal,
2005), mainly due to habitat transformation as
result of human activities, especially agricul-
tural expansion. Conservation of this species
requires a thorough understanding of its popu-
lation dynamics and phenological cycles, as
well as of the underlying factors.
Phenology studies the occurrence of repet-
itive biological events and the biotic and abi-
otic environmental factors that control them.
A better understanding of the phenology of
a plant species makes it possible to evaluate,
for instance, flower and fruit availability for
the local fauna (e.g., Morellato et al., 2000;
Cabrera & Wallace, 2007), or litter produc-
tion for soil formation and nutrient input (e.g.,
Vitousek, 1984; Sayer et al., 2020), all of
which are fundamental aspects to understand
forest dynamics.
Although the conservation and manage-
ment plan for C. quindiuense in Colombia
(Bernal et al., 2015) stresses the need to
conduct phenological studies, only a single
short-term work performed by Girón-Vander-
huck, Salazar, and Agudelo (2001a) is avail-
able. In this paper we present the results of a
two-year phenological study carried out along
an altitudinal gradient in the Tochecito River
basin, in the Central Cordillera of Colombia,
where the largest palm stands of this species
are found. We describe leaf production rate,
and flowering and fruiting cycles, and contrast
them among elevations. We also calculate the
food offer for the fauna, as a tool for the proper
management of this species in the area. As tem-
perature in the tropical Andes decreases by ca.
0.6º per 100 m increase in elevation (Ramírez,
Roldán, & Yépez, 2004), this change in tem-
perature is expected to influence phenology
along the gradient. Because no similar studies
appear to have been made in the Andes, we
expected differences in parameters to occur,
but did not have any clue of the direction of
variation, if any. We hypothesized that the
flowering sequence along the gradient would
somehow be similar to the sequential flower-
ing of several Ceroxylon species, described
by Carreño-Barrera, Madriñán, and Núñez-
Avellaneda (2013).
Study area: The study area is in the
Tochecito River basin, in the Department of
Tolima, on the Eastern slope of the Central
Cordillera of Colombia (4°30’45”-4°31’34”
N & 75°30’29”-75°26’31” W), between 2 400
and 3 000 m elevation (Fig. 1). It is a mountain
area of 8 890 ha, of which 43 % have some type
of palm cover (cloud forest remnants or sec-
ondary forests dominated by very dense palm
stands, cloud forest remnants or secondary
forests with some palms, matrix of pastures and
crops with very dense palm stands, and matrix
of pastures and crops with some palms) (Insti-
tuto Humboldt, 2016). The rainfall regime is
bimodal, with the rainiest period between June
and November, and the driest period between
February and March (IDEAM, 2020).
Revista de Biología Tropical, ISSN: 2215-2075, Vol. 69(2): 649-664, April-June 2021 (Published Jun. 09, 2021)
Study species: Ceroxylon quindiuense is
a single-stemmed palm, reaching up to 60 m in
height (Bernal et al., 2018) and 36 cm in diam-
eter at breast height. Its crown consists of 12-27
blue-green leaves, with the pinnae covered on
the underside by a woolly whitish indument.
It is a dioecious species, although, in some
cases, individuals change sex (Martínez, Sanín,
Castillo, López, & Bernal, 2018). The inflo-
rescences are interfoliar, repeatedly branched,
supported by a long peduncle, and bearing
several thousand small, whitish flowers, which
have anthesis almost simultaneously for one
to a few days (personal observation, 2012 to
2020; Carreño-Barrera et al., 2013). Although
the flowers of both sexes look very similar, sta-
minate and pistillate inflorescences are easily
recognized, even from a great distance. Pistil-
late inflorescences remain with their rachillae
completely extended, at a wide angle with the
rachis, even after fruiting and falling to the
ground. In the staminate inflorescences, on the
other hand, the rachillae are closely appressed
to the rachis after anthesis, so that the old inflo-
rescences have a broom-like appearance (Mar-
tínez et al., 2018). Each palm produces several
infructescences simultaneously and each one
bears thousands of spherical fruits, 1-2 cm in
diameter, smooth and orange-red when ripe
(Galeano & Bernal, 2010; Sanín & Galeano,
Data collection: We surveyed four sites
located at intervals of 200 meters along an
elevation gradient starting at 2 400 m.a.s.l. and
ending at 3 000 m.a.s.l., (2 400, 2 600, 2 800 and
3 000 m.a.s.l., Fig. 1). At each elevation, indi-
viduals were selected within a range of +/- 25
m with respect to the elevation value. The 2 400
m site was established in a secondary forest and
Fig. 1. Location of the Ceroxylon quindiuense study area in the Central Cordillera of Colombia. Sampling sites at different
altitudes: 1: Las Cruces (2 400 m.a.s.l.); 2: Galleguito (2 600 m.a.s.l.); 3-4: La Carbonera (2 800 and 3 000 m.a.s.l.). Based
on IGAC (2014) layers.