Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64854, mayo 2025 (Publicado May. 15, 2025)

Climate seasonality and plant vigor effects on reproductive

phenology of Potalia turbinata (Gentianaceae) in Costa Rica

Rachel Salazar1*; https://orcid.org/0009-0001-0275-5277

Jorge González-Linares2; https://orcid.org/0009-0002-7073-597X

Mauricio Fernández Otárola3; https://orcid.org/0000-0001-9240-7569

Gilbert Barrantes4; https://orcid.org/0000-0001-8402-1930

Bernal Rodríguez-Herrera5; https://orcid.org/0000-0001-8168-2442

1. Sistema de Estudios de Posgrado, Universidad de Costa Rica, San José, Costa Rica; rachel.salazar867@gmail.com

(*Correspondence)

2. Sistema de Estudios de Posgrado, Universidad de Costa Rica, San José, Costa Rica; jorge.gonzalezlinares@ucr.ac.cr

3. Escuela de Biología y Centro de Investigaciones en Biodiversidad y Ecología Tropical (CIBET), Universidad de

Costa Rica, San José, Costa Rica; mauricio.fernandez@ucr.ac.cr

4. Escuela de Biología y Centro de Investigaciones en Biodiversidad y Ecología Tropical (CIBET), Universidad de

Costa Rica, San José, Costa Rica; gilbert.barrantes@ucr.ac.cr

5. Escuela de Biología y Centro de Investigaciones en Biodiversidad y Ecología Tropical (CIBET), Universidad de

Costa Rica, San José, Costa Rica; bernal.rodriguez@ucr.ac.cr

Received 31-VIII-2024. Corrected 20-IV-2025. Accepted 21-IV-2025.

ABSTRACT

Introduction: Potalia turbinata (Gentianaceae) is a treelet distributed from Nicaragua to Panama. Information

on its reproductive biology and phenology is lacking.

Objective: To describe the reproductive phenology of P. turbinata, morphologically characterize its fruits and

seeds, and determine whether flower and fruit production correlates with climate (temperature and precipita-

tion) and plant vigor.

Methods: The research took place at Tirimbina Biological Reserve in Costa Rica, January 2017–March 2019.

Monthly counts of flowers and fruits were conducted on 86 plants. We used circular statistics to determine the

duration and peak of flowering and fruiting. Fruit production was compared between years to describe repro-

ductive cycles. We correlated flower and fruit abundance with climatic conditions. Plant measurements included

height, basal stem diameter, leaf crop, seeds per fruit, fruit and seed dimensions, and fruit hardness. We used

height, diameter, and number of leaves as proxies for plant vigor and compared plant vigor between fruiting (i.e.,

number of fruits) and non-fruiting plants.

Results: Flowering lasted 2–6 months per plant, peaking in April, while fruiting was prolonged (3–10 months),

moderately peaking in August. Flower abundance was negatively correlated with precipitation, but fruiting

showed no significant correlation with climate. Fruit production negatively correlated with plant height and

positively correlated with basal stem diameter. Non-fruiting plants had more leaves than fruiting plants.

Conclusions: At the population level, P. turbinata flowered and fruited annually. Individuals were supra-annual,

an uncommon reproductive pattern for tropical species. The correlation between flowering and lower precipita-

tion supports possible insect pollination, though direct pollinator observations remain needed. The results con-

cur with other studies indicating a correlation between fruit number and plant size. This investigation provides

information on the reproductive phenology and fruit traits of P. turbinata, laying a foundation for future research

into its interactions with pollinators and dispersers.

Keywords: flower and fruit production; Gentianaceae; treelet phenology; tropical forest; fruit morphology;

Tirimbina Biological Reserve.

https://doi.org/10.15517/rev.biol.trop..v73iS2.64854

SUPPLEMENT

SECTION: REPRODUCTION

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64854, mayo 2025 (Publicado May. 15, 2025)

INTRODUCTION

Phenology refers to the study of life cycle

events and the seasonal timing of each event for

individuals, populations, species, and commu-

nities (Rathcke & Lacey, 1985). Plant reproduc-

tive phenology includes the timing and patterns

of flowering and fruiting (Engel & Martins,

2005; Boyle & Bronstein, 2012; Numata et al.,

2022). Reproductive phenological events are

influenced by abiotic factors such as precipita-

tion and temperature, ecological interactions

with pollinators and dispersers (Pires et al.,

2018), as well as phylogenetic and genetic

constraints (Wright & Calderon, 1995; Fenner,

1998; Wilczek et al., 2010; Du et al., 2020).

Investigating the phenological strategies and

life cycles of plants provides insights into their

reproduction and survival (Rathcke & Lacey,

1985) as well as their interactions with herbi-

vores and other animals which use the plants

(van Schaik et al., 1993). Detailed information

on reproductive phenology is still lacking for

many tropical plant species.

Phenological patterns are more variable in

tropical wet forests than temperate and tropical

dry forests as the environment supports flower-

ing and fruiting year-round (Opler et al., 1980;

Boyle & Bronstein, 2012). Variation in tropical

plant phenology is reflected in their diverse pat-

terns that include subannual (reproducing more

than once per year), annual (reproducing once

a year), continuous (flowering and fruiting all

year), and supra-annual cycles (one reproduc-

tive cycle over more than one-year) (Newstrom

et al., 1994). Temperate plant species are more

RESUMEN

Efectos de la estacionalidad climática y el vigor de las plantas sobre la fenología reproductiva

de Potalia turbinata (Gentianaceae) en Costa Rica

Introducción: Potalia turbinata (Gentianaceae) es un arbusto distribuido desde Nicaragua hasta Panamá. La

información sobre la biología de esta especie es escasa y se desconoce su fenología reproductiva.

Objetivo: Describir en detalle la fenología reproductiva de P. turbinata, caracterizar morfológicamente sus frutos

y semillas, y determinar si la producción de frutos y flores se correlaciona con el clima (temperatura y precipit-

ación) y el vigor de la planta.

Métodos: La investigación se llevó a cabo en la Reserva Biológica Tirimbina, Sarapiquí, Costa Rica, desde enero

2017 hasta marzo 2019. Se realizaron conteos mensuales de flores y frutos en 86 plantas. Se utilizaron estadísticas

circulares para determinar la duración y el pico de las fenofases de flores y frutos. La producción de frutos se com-

paró entre años para describir los ciclos reproductivos. Se probó la correlación entre la cantidad de flores y frutos

y la estacionalidad climática. Las mediciones de las plantas incluyeron altura, diámetro basal del tallo, número de

hojas, semillas promedio por fruto, dimensiones de frutos y semillas, y dureza de los frutos. El vigor de la planta

(altura, diámetro basal, número de hojas fueron variables usadas como una aproximación de vigor) se evaluó en

relación con el tamaño de la cosecha de frutos.

Resultados: La floración duró entre 2 y 6 meses por planta, con un pico en abril, mientras que la fructificación

fue prolongada (3–10 meses), con un pico moderado en agosto. La abundancia de flores se correlacionó negativa-

mente con la precipitación, pero la fructificación no mostró una correlación significativa con el clima. La produc-

ción de frutos se correlacionó negativamente con la altura de la planta y positivamente con el diámetro basal del

tallo. Las plantas sin frutos tenían más hojas que las plantas con frutos.

Conclusiones: A nivel de población P. turbinata tiene un ciclo reproductivo anual pero los individuos tienen una

reproducción irregular supra-anual, un ciclo reproductivo poco común en especies tropicales.

La correlación entre la floración y menor precipitación respalda la posibilidad de polinización por insectos,

aunque aún se requieren observaciones directas de polinizadores. Los resultados coinciden con otros estudios que

indican una correlación entre el número de frutos y el tamaño de la planta. Esta investigación aporta información

sobre la fenología reproductiva y las características de los frutos de P. turbinata, sentando las bases para futuras

investigaciones sobre sus interacciones con polinizadores y dispersores.

Palabras clave: producción de flores y frutos; Gentianaceae; fenología de arbustos; bosque tropical; morfología

de frutos; Reserva Biológica Tirimbina.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64854, mayo 2025 (Publicado May. 15, 2025)

seasonal and predictable than tropical species

and often closely follow temperature, precipita-

tion, and light diel cycles (Ting et al., 2008).

In a tropical dry forest, where water stress

influences reproductive phenology, tree species

tend to flower in response to seasonal rainfall,

with dry-season flowering linked to drought-

resistant species (Lasky et al., 2016).

Precipitation influences productivity and

seasonality in tropical environments (Brearley

et al., 2007). Many studies report that fruiting

intensity and peaks increase with precipita-

tion in tropical rainforests (Mendoza et al.,

2017; Dunham et al., 2018), and in Costa Rica,

understory species show a fruiting peak in

the second half of the year during the rainier

period (Opler et al., 1980; Boyle & Bronstein,

2012). Understory species have irregular flow-

ering in rain forests, but more commonly,

shrubs show higher flower production during

the less rainy period at the beginning of the year

(Opler et al., 1980).

Plant flowering and fruiting are often

influenced by plant size because larger plants

can allocate more resources for reproduction

(Ollerton & Lack, 1998; Fernández Otárola et

al., 2013), and investigating the relationship

between plant size and fruit production pro-

vides insight about the strategies implemented

by the plant (e.g., resource allocation and size

at reproductive maturity). Larger plants tend

to have more and/or larger fruits (Herrera,

1993). A study on the legume Lotus cornicula-

tus showed that larger plants produced more

fruits and had lower seed predation (Ollerton

& Lack, 1998). In a wet tropical forest of Costa

Rica, 17 tree species showed the same pattern,

with larger plants producing more fruits; tree

size also predicted whether a plant reproduced

(Minor & Kobe, 2019). A study in a lowland

conifer forest investigated the effect of plant

size, plant age, site factors, and canopy density

on flower and fruit production of nine under-

story shrubs; the study stated that plant size

(closely followed by plant age) was the main

factor in predicting flower and fruit quantities

(Wender et al., 2004).

The family Gentianaceae comprises 16

genera and 30 hermaphroditic species in Costa

Rica (Sánchez, 2010). These species predomi-

nantly consist of herbs, although there are some

shrubs, trees, mycoheterotrophs, and epiphytes

(Sánchez, 2010). Potalia turbinata Struwe & V.

A. Albert is an understory treelet and the only

species of the genus in Costa Rica. Flowers of

P. turbinata have been observed from March

to May, and fruits in May, July, September, and

December (Struwe & Albert, 2004). There is

currently no formal description of the phenol-

ogy of this plant or the other eight species in the

genus (Frasier et al., 2008).

Information on the reproductive phenol-

ogy of P. turbinata would facilitate the study of

their use by floral visitors and frugivorous spe-

cies. Potalia turbinata is known to be utilized by

bats; for instance, Vampyriscus nymphaea uses

its leaves to build conical tents as refuge and

feeding sites (Villalobos-Chaves et al., 2016).

Tent construction may reduce flower produc-

tion by limiting the foliar area exposed to the

low light in the forest understory (Rodríguez-

Herrera et al., 2018). Describing the phenology

of P. turbinata establishes a foundation for fur-

ther studies on the dynamics of its interactions

with associated fauna.

The aim of this study was: 1) to describe

the reproductive phenology of P. turbinata and

the morphological characteristics of its fruits

and seeds, 2) to determine whether rainfall and

temperature correlate with flower and fruit pro-

duction in the population, and 3) to evaluate if

plant vigor (height, basal stem diameter, and

number of leaves) correlates with fruit produc-

tion. We expected P. turbinata to adjust its phe-

nology to maximize reproductive success. Since

this species is likely insect-pollinated (Rincón

et al., 1999), we predict that it will flower during

the early months of the year when precipitation

decreases and conditions are more favorable

for insect pollinators (Janzen, 1967). On the

contrary, we expected that fruiting occur with

the onset of heavier rains in the latter half of

the year, as happens in many insect-pollinated

plants in rainforests. Temperature exhibits min-

imal variation throughout the year and should

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64854, mayo 2025 (Publicado May. 15, 2025)

not correlate with either of the phenophases

(flowering and fruiting). We expected plant

vigor to correlate with fruit production.

MATERIALS AND METHODS

Study site and species: We conducted

this study at the Tirimbina Biological Reserve

(hereafter Tirimbina), Heredia province, Costa

Rica (10°25’ N; 84°47’ W). Tirimbina includes

345 ha mostly covered by mature rain forest

(Holdridge et al., 1971). The mean annual tem-

perature and precipitation are 24.3 °C and 3 777

mm respectively, with an elevation of 180–220

m asl (Ley-López & Avalos, 2017). The rainy

season occurs from May through November,

while the dry season extends from December

through April.

Potalia turbinata is endemic to Central

America and has a restricted and patchy distri-

bution in southeastern Nicaragua, northeastern

Costa Rica, and northeastern Panama (Struwe

& Albert, 2004; Sánchez, 2010). Its habitat

ranges from sea level to 700 m (Sánchez, 2010).

This species is a pachycaul understory shrub

or treelet with a height up to 4.5 m. The leaves

are simple, 30–75 cm long and 7.5–18 cm wide,

oblanceolate, and grouped distally on branches

(Sánchez, 2010). The flowers are produced on

a compact corymbose-cymose inflorescence,

with up to 30 flowers that vary in color from

white, yellow, pale green, yellow-green, or green

(Struwe & Albert, 2004; Sánchez, 2010). The

green fruits are turbinate with a transverse

ring and may darken slightly as they mature

(Sánchez, 2010).

Phenology: We located, marked, and

quantified P. turbinata plants in a high-den-

sity area along the Ajillo trail (ca. 805 m

long), including all individuals within 10 m on

either side of the trail. We sampled monthly

from January 2017 to March 2019, covering a

total of 27 months. A total of 83 P. turbinata

plants were sampled in 2017, increasing to 86

in 2018 and at the beginning of 2019. Sam-

pling frequency may affect the interpretation

of phenological patterns. For tropical trees, it

is advised to sample every two weeks, but if

monthly sampling is chosen, a minimum of 25

trees are recommended (Morellato et al., 2010).

Fournier and Charpantier (1975) comment

that monthly observations are appropriate to

obtain a good representation of tree phenology;

particularly in species that phenological phases

change slowly as is the case of Potalia turbinata

that changes slowly from flowering to fruit-

ing and fruit maturation (Rodríguez-Herrera,

unpubl.). Hence, we considered monthly sam-

pling appropriate.

During each survey, we counted flower

buds, flowers in anthesis, and fruits for each

plant, rather than per inflorescence. The num-

ber of inflorescences was not recorded. Imma-

ture and mature fruits were grouped together

due to difficulty in distinguishing their devel-

opment conditions because of the lack of con-

spicuous changes in color or size. To classify the

different reproductive stages of the plant, the

reproductive structures were classified as flow-

er buds if plants had unopened flowers, flower-

ing if at least one flower was open, and fruiting

if they had fruits at any stage of development.

A plant could have more than one reproductive

stage at a time.

The CircStat package (Berens, 2009) was

used to determine the circular mean and the

value of the vector-r for each phenophase.

Vector-r varies between 0 and 1, where values

closer to 1 indicate a strong seasonality (i.e., a

defined peak), and values closer to 0 indicate

low or absence of seasonality. To determine

if the circular distribution was seasonal, we

applied the Rayleigh test (Zar, 1999). This

information facilitated the determination of

the duration, peak, and frequency of the flower

and fruit phenophases at the population level.

Data from 83 plants from 2017 and 2018 were

combined in this analysis.

We implemented a Student’s t-test to com-

pare fruit production between 2017 and 2018

to assess interannual variability as basal infor-

mation on the annual variation in resources

for animals. Fruit production data were log-

transformed to improve normality. Normality

was assessed using the Shapiro-Wilk test, and

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64854, mayo 2025 (Publicado May. 15, 2025)

variance homogeneity was tested using Lev-

ene’s test. Fruit production is defined as the

maximum number of fruits observed on each

plant at a given census rather than cumulative

fruit output over the entire reproductive season.

This method was chosen because cumulative

fruit counts were not feasible in our monthly

sampling design, and the maximum observed

count provides a standardized comparison

across individuals and years.

We tested for correlation (Pearson’s cor-

relation) between monthly 24-hour average

precipitation and temperature with monthly

total numbers of flowers and fruits in the popu-

lation during the study period. To account for

potential time-lagged effects of climate season-

ality on the reproductive phenology, we tested

for correlations with 0-, 1-, 2-, and 3-month

offsets of meteorological data. Climate data

with records for the duration of the study were

obtained from a local weather station at Tir-

imbina. This analysis helps determine whether

climatic factors such as rainfall or temperature

act as cues for triggering flowering and fruiting,

potentially influencing pollination success and

seed dispersal.

Plant morphological traits: We recorded

the morphological measurements for 81 plants

in January 2019. We measured the height of

each plant from the ground to the top of the

apical meristem, basal stem diameter at 10

cm above ground, and recorded the num-

ber of fully developed leaves per plant. Three

plants had modified leaves in 2018, presumably

altered by tent-roosting bats. These leaves were

included in the study as they were still living

and attached to the plant. Additionally, these

plants had already initiated their reproductive

cycles before leaves were modified.

In addition to the plant vigor traits, we

measured fruit and seed characteristics. Fruits

with well-developed transverse rings were col-

lected from plants near the study area to mea-

sure their morphological characteristics (2–10

fruits per plant), such as weight (OHAUS Scout

Pro scale, model SPE 123), length, and width at

the widest point of each fruit’s transverse ring.

A total of 185 fruits from 27 plants were col-

lected (19 in August 2017, 106 in October 2017,

and 60 in November 2018), and fruit hardness

was determined for 132 fruits. Fruit hardness,

defined as the strength required to perforate

the exocarp and mesocarp and access the seeds,

was estimated using a penetrometer with a 2

mm chisel (Chatillon, model DPPH-100). The

chisel was aligned perpendicular to the fruits’

transverse rings. We weighed 150 seeds from

30 fruits collected from 10 plants, with five

seeds taken per fruit, using a precision lab scale

(Sartorius, model GMBH 2842), and measured

their length, width, and thickness. Fruit and

seed dimensions were measured with a dial

caliper (SPI, model 31-414-6).

Plant vigor effects on fruit production:

To evaluate the effect of plant vigor on fruit

production of P. turbinata, we used a general

linear model with fruit production per plant

as the response variable and height, basal stem

diameter, and number of leaves as explana-

tory variables. The explanatory variables were

standardized (mean-centered and scaled by

standard deviation) to allow for interpreta-

tion of model coefficients on a common scale.

This analysis focused on data from the 2018

reproductive season because vigor traits from

the previous year were incomplete. Addition-

ally, we tested whether plant vigor traits dif-

fered between fruiting and non-fruiting plants

using a general linear model, with each trait

as a response variable and reproductive status

(fruiting vs. non-fruiting) as the explanatory

variable. Since a minimum reproductive size

has not been formally established for P. t u r bi -

nata, no size threshold was applied to exclude

potential juveniles among the non-fruiting

plants. Additionally, individuals at the mini-

mum recorded height and basal stem diameter

in our dataset were observed fruiting. All statis-

tical analyses were conducted in R v.4.0 (R Core

Team, 2024).

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64854, mayo 2025 (Publicado May. 15, 2025)

RESULTS

Phenology: Flower buds began in Febru-

ary, with a peak in March (Fig. 1A). Records

of open flowers began in March and peaked in

April (Fig. 1B), about a month after the flower-

bud peak. Fruit production was prolonged,

starting in April and continuing until February

of the following year, with peak production

occurring in May, July, and August (Fig. 1C);

June 2018 was not sampled, and this may have

affected the fruiting pattern observed in Fig.

1C. Flower bud and flower production were

highly seasonal, with a significant unimodal

pattern (Fig. 1A, Fig. 1B). However, fruit pres-

ence showed a more prolonged, barely seasonal

pattern (Fig. 1C). The onset of the different

floral stages did not vary between the two sam-

pling years.

The presence of fruits lasted for up to 10

months from the initial detection of the fruit-

ing phase until all fruits had either fallen from

the plants or been removed. The fruiting period

of individual plants averaged 7 months with a

range of 3 to 10 months. Flower bud and flower

presence per plant ranged from 2 to 6 months.

The number of plants flowering each year

varied up to 369%. There were 13 flowering

plants in 2017, 48 in 2018, and 20 by March

2019. All plants that flowered in 2017 and 2018

produced fruits; observations in 2019 ended

before the fruiting phase. Nine plants (69.2 %)

flowered subsequently from 2017 to 2018, and

18 plants (37.5 %) from 2018 to March 2019.

Four plants that produced flowers in 2017 also

produced flower buds in 2019, three of which

flowered all three years.

Fruit production was significantly lower in

2017 than 2018 (28.15 ± 21.24 SD versus 43±

27.16 SD: t = -2.07, df = 56, p = 0.043). The

number of fruits per plant at the peak count

ranged between 10 to 81 in 2017, and 4 to 136

fruits in 2018. The mean value of the peak count

of fruits per fruiting plant during the study was

39.67 (± 26.53 SD) and for flowers it was 29.94

(± 21.81 SD). The number of flowers observed

per plant ranged from 2 to 107. The maximum

fruit count across the population was 344 fruits

in June 2017 and 1667 in May 2018. Flower

buds and flowers considered together had a

maximum count across the population of 543

in 2017 and 1658 in 2018, both in March.

The number of flowers in the population

correlated negatively with monthly precipita-

tion (r=-0.442, p=0.040), so the number of

flowers increased when precipitation decreased.

However, the number of fruits did not correlate

with precipitation (r=0.359, p=0.101). Flower-

ing remained negatively correlated with precip-

itation at a 1-month offset (r=-0.451, p=0.040),

but this relationship weakened at 2-month (r=-

0.087, p=0.709) and 3-month offsets (r=-0.046,

p=0.842). Fruit production did not correlate

significantly with precipitation at any offset

Fig. 1. Rose graphs of reproductive phenology in Potalia turbinata (Gentianaceae) showing the percentage of individuals

with: A. flower buds, B. open flowers, and C. fruits. The arrow indicates the circular mean, and its length represents the value

(0-1) of the vector-r (greater length equals greater seasonality). P-values for the Rayleigh test indicate a clear seasonality of

the three reproductive stages.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64854, mayo 2025 (Publicado May. 15, 2025)

(all p>0.1). Flowering (r=0.017, p=0.939) and

fruiting (r=-0.170, p=0.449) did not correlate

with temperature which had a 24-hour aver-

age monthly temperature of 25.5 °C, with a

narrow range from 22.4 to 31.7 °C during the

study. Correlations with temperature were non-

significant across all offsets (all p-values>0.1).

Plant morphological traits: Table 1 sum-

marizes the measurements of the plants, fruits,

and seeds. Of the 81 plants measured in January

2019, 55.6% had fruited in 2018.

Plant vigor effects on fruit production:

Among the analyzed plant vigor traits, basal

stem diameter showed a strong positive correla-

tion with fruit production (Table 2). In contrast,

taller plants were associated with lower fruit

production (Table 2; Fig. 2). Of these variables,

only the number of leaves was significantly

Table 1

Descriptive values of plant vigor, fruit, and seed traits of Potalia turbinata (Gentianaceae).

Plant vigor traits Height (cm) Basal Stem Diameter (cm) Number of Leaves

Mean ± SD 245.7 ± 80.4 3.2 ± 0.8 16.9 ± 6.2

Range 60–450 1.9-5.1 4–38

Fruit traits Weight (g) Length (mm) Width (mm) Hardness (N)

Mean ± SD 2.7 ± 0.5 18.9 ± 1.1 21.3 ± 1.6 44.6 ± 8.1

Range 1.4–3.9 16.0–21.5 16.1–24.7 16.0–68.0

Seed traits Seeds per Fruit Weight (mg) Length (mm) Width (mm) Thickness (mm)

Mean ± SD 36.2 ± 7.3 13.8 ± 2.8 4.8 ± 0.4 2.9 ± 0.3 1.4 ± 0.2

Range 11.0–46.0 1.3–20.4 3.6–5.6 1.8–3.8 0.8–1.9

Table 2

Results of a GLM on the effect of plant vigor traits on fruit production in Potalia turbinata (Gentianaceae).

Factor Estimate S.E. t-value P-value

Intercept 42.91 2.95 14.53 <0.001

Height -12.51 4.24 -2.95 0.005

Diameter 26.24 4.23 6.21 <0.001

No. leaves 4.51 3.00 1.50 0.140

Fig. 2. The relationship between A. plant height, B. basal stem diameter, and C. number of leaves with fruit production in

plants of Potalia turbinata (Gentianaceae) in Costa Rica.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64854, mayo 2025 (Publicado May. 15, 2025)

higher in non-fruiting plants (p=0.007); the

other two variables did not differ between fruit-

ing and non-fruiting plants (Fig. 3).

DISCUSSION

Phenology: Phenological patterns in flow-

ering and fruiting often differ between indi-

vidual and population levels (Sakai et al., 2005;

Fenner, 1998). In contrast, a study at La Selva

(20 km from Tirimbina) analyzing 226 treelets

and shrubs across 35 species found that despite

some individual variation, phenological pat-

terns at the individual level generally reflected

those of the population (Boyle & Bronstein,

2012). Potalia turbinata exhibited lower con-

sistency between individual and population-

level phenology. As a population, P. turbinata

followed an annual phenological pattern, but

differed at the individual level with a small

fraction of the individuals exhibiting annual

patterns, while the majority were supra-annual.

Many species follow an annual cycle at La

Selva in accordance with precipitation fluc-

tuation, similar to the documented pattern in

Potalia turbinata at the population level (Sakai,

2001). At the individual level at La Selva, suban-

nual phenological patterns were predominant,

followed by annual patterns, while continuous

and supra-annual strategies were rare (News-

trom et al., 1994; Sakai et al., 2005), making P.

turbinata a notable exception at the individual

level. This suggests that P. turbinata individuals

may require longer recovery periods between

reproductive events, potentially due to resource

limitations or other ecological constraints.

Several factors influence fruit and seed

production, including resource allocation, floral

herbivory, and effective pollination (Cunning-

ham, 2000). Insufficient resource accumulation

in certain plant species can lead to a supra-

annual fruiting pattern, as these plants require

longer periods to replenish between fruiting

years (Fenner, 1998). Boyle and Bronstein

(2012) mentioned light variation, rainfall, and

nutrient availability as possible factors affecting

flowering and fruiting patterns; although, in

their study, precipitation was not an influencing

factor. Other contributing factors to variation

include competition and herbivory (Sakai et al.,

2005). Cunningham (2000) studied an under-

story plant and found that floral herbivory was

the main limiting factor on fruit production

followed by the proportion of flowers visited by

pollinators. Genetics may also influence plant

phenology, flowering, and fruiting (Wilczek et

al., 2010; Tang, et al., 2016). Abiotic and biotic

factors may control plants with conservative

genomes to a lesser extent (Fenner, 1998).

The average annual fruit production in

Potalia turbinata was significantly lower in

2017 than in 2018, highlighting interannual

variation in reproductive output. This varia-

tion further suggests that resource availability

and environmental factors may have a stron-

ger influence on fruit production than floral

Fig. 3. Comparison of plant vigor traits between fruiting plants and non-fruiting plants of Potalia turbinata (Gentianaceae)

in Costa Rica. Data from the year 2018: A. height, B. basal stem diameter, and C. number of leaves.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64854, mayo 2025 (Publicado May. 15, 2025)

herbivory, which was not observed during this

study. While we described flowering and fruit-

ing patterns, future research could examine

additional reproductive metrics, such as fruit

set and seed viability, to better understand

reproductive success in P. turbinata.

Tropical plants tend to have longer fruit-

ing periods, averaging four months (Jordano,

1992). Potalia turbinata is one example of an

extremely lengthy fruiting period at the popula-

tion level (ten months) and at the level of indi-

vidual plants (approximately seven months).

In this study, immature and mature fruits were

considered together due to difficulty in dis-

tinguishing the categories because of cryptic

color and size changes. Immature fruits in early

development present an underdeveloped trans-

verse ring. In future studies, to avoid ambiguity

while deciding whether fruits are immature or

mature, color could be measured and the diam-

eter of the fruit ring across the months could be

considered. Though it would be more demand-

ing in the field, a higher resolution of the timing

of phenological events could be achieved, such

as a more precise timing of fruit maturation.

Additionally, the start of fruit dispersal could be

used as an indicator of fruit maturity.

Potalia turbinata only had one flower-

ing period per year at Tirimbina. Flowering

took place at the beginning of the year (lower

precipitation) and fruiting at the middle to

second half of the year (higher precipitation),

which is consistent with flowering and fruit-

ing tendencies reported by Opler et al. (1980)

of treelets and shrubs in Costa Rican tropical

wet forests. Flowering periods were shorter for

supra-annual species and tended to occur in the

dry season more than for annual or subannual

plants (Bawa et al., 2003), which is similar to

P. turbinata with a population level flowering

period of four to five months during a period

of less rainfall.

Flowering in the dry season may influence

pollination success in P. turbinata by aligning

reproductive timing with seasonal changes in

pollinator availability. Tree and shrub species

pollinated by bees, butterflies, flies, and wasps

tend to reach their peak flowering during the

dry season in Venezuela (Ramírez, 2006). All

flowering plants of P. turbinata successful-

ly produced fruits and exhibited pronounced

flowering seasonality. Plants with high flower-

ing synchrony are more likely to reproduce

successfully, as they attract more pollinators

(Bruno et al., 2019). Successful cross-pollina-

tion, autonomous self-pollination, and apomix-

is are possible mechanisms contributing to seed

development. While P. turbinata had strong

flowering seasonality, its pollination strategy

remains uncertain. Rincón et al. (1999) sug-

gested that the species may be insect-pollinated,

but no direct observations confirm this. Fur-

ther research is needed to identify pollination

strategies and explore the environmental fac-

tors influencing fruit production in P. turbinata.

Though the onset of the rainy season

was not a significant determinant in the P.

turbinata fruiting cycle, the flowering signifi-

cantly correlated with less rain which in turn

led to the presence of fruit during the rainy

months. Seed germination facilitation is one

factor that may influence fruiting occurring

during the rainy season (Garwood, 1983). The

lack of correlation between temperature and

flowering and fruiting may be a consequence

of the little, non-seasonal variation in this

environmental variable.

The timing of flowering and fruiting of P.

turbinata in this study overlapped with herbar-

ium collected phenological information. Sán-

chez (2010) states that flowers are found from

March to June, while Struwe and Albert (2004)

mention March to May and specify fruiting

observations in May, July, September, and

December. This investigation further detailed

the reproductive phenophases by examining

their full extension and patterns.

Plant morphological traits: The plant

height of P. turbinata plants in this study is con-

sistent with the values reported by Struwe and

Albert (2004). The number of flowers was pre-

viously reported as up to 30 per plant (Struwe &

Albert, 2004), but we observed a range beyond

that number. We also documented a larger stem

diameter at 3.21 cm, compared to 14 mm, and

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64854, mayo 2025 (Publicado May. 15, 2025)

fruit size was slightly larger in this study (16–22

x 16–25 mm compared to 10–14 x 11–18 mm).

However, Struwe and Albert (2004) collected

measurements from dry material which could

lose 10–15 % of its mass. Sánchez (2010)

described the seeds as 4–5.5 mm which is in

accordance with our findings. Additionally, we

provided novel data on the number of seeds,

weight, width, and thickness, as well as fruit

hardness.

Plant and fruit morphological traits are

noteworthy for further investigations involving

fruit consumption or seed dispersal mecha-

nisms of P. turbinata. The seeds of Neotropi-

cal woody species are primarily dispersed by

vertebrates (Howe & Smallwood, 1982), and

the size of the seeds influences whether they

are ingested (Sebastián-González, 2017, Ong et

al., 2021). Smaller seeds are more likely to be

swallowed by a disperser and pass through the

digestive tract; larger seeds (> 12 mm) tend to

only be swallowed by large vertebrates includ-

ing primates, tapirs, and peccaries (Fuzessy et

al., 2018). Seed weight and length can influence

whether a seed is swallowed or dropped as well

as which frugivores act as dispersers. Large-

seeded fruits are less likely to be consumed by

birds, bats, and small mammals (Fuzessy et al.,

2018), although bats and small sized rodents

have been observed dispersing large-seeded

fruits by synzoochory (Melo et al., 2009; Ong et

al., 2021). Potalia seeds are small and numerous,

suggesting they are likely dispersed through gut

passage, as observed in Carollia perspicillata

dispersing Potalia amara seeds (Lobova et al.,

2009). Bats have also been observed removing

fruits of P. turbinata (Salazar, unpubl.). Further

investigation into Potalia seed dispersal is war-

ranted due to a lack of information.

Fruit hardness may also affect fruit con-

sumption as there is a limit to the hardness that

can be handled by different species. Dumont

(1999) documents changing behaviors of phyl-

lostomid bats when they consume fruits of

varying hardness and indicates that some spe-

cies have morphological traits that aid in con-

sumption. Additional studies on P. turbinata

could contribute information on animal diets,

particularly regarding fruit hardness.

Plant vigor effects: The height of fruiting

plants of P. turbinata was negatively correlated

with fruit production, while basal stem diam-

eter showed a positive correlation with fruit

production. These findings partially align with

previous studies reporting that tree size is often

positively associated with reproductive out-

put or influences initial reproductive activity

(Herrera, 1991; Susko & Lovett-Doust, 2000;

Fernández-Otárola et al., 2013). A study at La

Selva Biological Station of approximately 2,000

trees from 17 species showed that the number

of fruits produced could be predicted by plant

size (Minor & Kobe, 2019). This trend has also

been shown in other plants such as a cactus

species studied by Bustamante and Búrquez

(2008), which had higher flowering intensity

and plant fecundity in larger plants.

The amount of light available through

canopy gaps may influence plant size (Wender

et al., 2004). Competition can also affect plant

size, and increased plant growth usually cor-

relates with more access to resources; crowd-

ing affects the plant reaching its maximum

potential size and in turn, affects its crop size

or fecundity (Tracey & Aarssen, 2011). Fig

tree fruit production may be influenced by the

amount of time the trees are able to restore

nutrients between reproductive cycles (Huang

et al., 2019), which is another factor along with

plant size that could be considered in future

studies on P. turbinata.

The number of leaves did not correlate

with fruit production in P. turbinata when ana-

lyzing only fruiting plants, indicating that leaf

number does not directly influence the number

of fruits. However, there was a significant dif-

ference in leaf number between fruiting and

non-fruiting plants, with non-fruiting plants

having more leaves, while fruiting plants had

fewer. This pattern suggests an inverse rela-

tionship between leaf production and fruiting,

potentially driven by resource allocation trade-

offs. This finding aligns with studies show-

ing a balance between vegetative growth and

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64854, mayo 2025 (Publicado May. 15, 2025)

reproductive effort. Wheelwright (1986) found

that fruit production and new leaf production

in the same year were negatively correlated in

12 of 15 studied tree species of the Lauraceae

family in Monteverde, Costa Rica. In most spe-

cies, vegetative growth and reproduction were

negatively associated, which may explain why P.

turbinata fruiting plants exhibited fewer leaves.

This suggests that resource allocation toward

fruit production may come at the expense of

vegetative growth. In trees, vegetative growth

(leaf production) and reproductive develop-

ment (flower and fruit production) are closely

linked (Borchert, 1983). The extent and driv-

ers of this potential trade-off in P. turbinata

remain unclear.

Previously, specific phenological informa-

tion was lacking for the genus Potalia apart

from knowing in which months flowering and

fruiting herbarium specimens were collected.

We found that P. turbinata produces flowers

and fruits annually at the population level with

varying patterns at the individual level. Higher

flower abundance coincides with drier months,

as predicted. The initial expectation of the

fruiting phenophase correlating with rainfall

was not supported. Fruit production of P. t u r -

binata correlates with plant size (height and

diameter), concurring with predictions. The

information gathered serves as a foundation

for further investigations on P. turbinata. Next

steps include identifying pollinator and dis-

perser interactions with the plant and the abiot-

ic factors that influence phenological strategies.

Ethical statement: The authors declare

that they all agree with this publication and

made significant contributions; that there is no

conflict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

We thank Emmanuel Rojas Valerio for his

help during the field work and Marco Vinicio

Sáenz Murillo at the Centro de Investigaciones

Agronómicas (CIA) at Universidad de Costa

Rica for his assistance in measuring the fruits

and seeds of the plant of study. Furthermore,

we would like to express our appreciation to the

staff at the Tirimbina Biological Reserve.

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