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Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64533, mayo 2025 (Publicado May. 15, 2025)
Effect of elevation, habitat, and season in butterfly
(Lepidoptera: Papilionoidea) assemblages in a tropical mountain
Wendolyn Matamoros-Calderón1, 2; https://orcid.org/0000-0002-2715-1875
Ricardo Murillo Hiller1,4; https://orcid.org/0000-0001-5666-8446
Álvaro Cerdas Cedeño3; https://orcid.org/0009-0000-6005-886X
Paul Hanson Snortum1, 3; https://orcid.org/0000-0002-7667-7718
Noemi Canet Moya3; https://orcid.org/0009-0003-5015-8519
Eduardo Chacón-Madrigal2, 5 *, https://orcid.org/0000-0002-8328-5456
1. Museo de Zoología, Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa
Rica, San Pedro de Montes de Oca, San José, Costa Rica; wendolyn.matamoros@ucr.ac.cr, murillohiller@gmail.com,
phanson91@gmail.com
2. Herbario Luis Fournier Origgi, Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de
Costa Rica, San Pedro de Montes de Oca, San José, Costa Rica; edchacon@gmail.com (*Correspondencia).
3 Escuela de Biología, Universidad de Costa Rica, San Pedro de Montes de Oca, San José, Costa Rica; alvaro.cerdascede-
no@ucr.ac.cr, nmcanet@gmail.com
4. Licenciatura en Gestión Ecoturística, Sede de Guanacaste, Universidad de Costa Rica,Liberia, Guanacaste, Costa Rica.
5. Herbario Nacional, Departamento de Historia Natural, Museo Nacional de Costa Rica.
Received 28-VIII-2024. Corrected 04-III-2025. Accepted 07-III-2025.
ABSTRACT
Introduction: Spatial and temporal variation in the environment promotes biological diversity. However, the
combined effects of elevation, seasonality, and habitat on butterfly diversity and abundance are still underex-
plored in rural tropical ecosystems. Butterflies, as bioindicators, offer an excellent opportunity to assess environ-
mental impacts due to their sensitivity to habitat changes.
Objective: This study aimed to evaluate the variation in butterfly abundance and species richness across different
habitats, elevations, and seasons in a rural tropical mountain landscape in Costa Rica.
Methods: The study was conducted over six months, encompassing both the dry and rainy seasons. Butterflies
were sampled along an elevational gradient (1 200, 1 500, and 1 800 m) across two habitat types: grasslands and
riparian forests. Sampling methods included hand nets and fruit baits along each transect and at each site over
the six-month period. We made six visits to each site, with seven transects sampled per visit to capture butterflies.
A total of 1 421 individuals representing 151 species were recorded.
Results: The Nymphalidae family exhibited the highest species richness and abundance. Butterfly abundance
decreased with increasing elevation, but patterns of species richness varied by family. Seasonal variation sig-
nificantly affected both abundance and richness, with higher values recorded during the rainy season. Based on
local abundance, rare species were more commonly found in riparian forests at higher elevations during the wet
season. Habitat did not significantly influence overall butterfly abundance or richness.
Conclusions: Seasonality and elevation influence butterfly populations in tropical mountain ecosystems. The
lack of a significant habitat effect suggests that other ecological variables may mediate the impact of habitat on
butterfly communities. Further studies are required to clarify these dynamics, particularly the role of habitat
heterogeneity.
Keywords: elevational gradient; landscape ecology; biodiversity patterns; population ecology; tropical forest.
https://doi.org/10.15517/rev.biol.trop..v73iS2.64533
SUPPLEMENT
SECTION: ECOLOGY
2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64533, mayo 2025 (Publicado May. 15, 2025)
INTRODUCTION
Biological diversity is crucial for ecosys-
tem functioning (Mace et al., 2012) and varies
spatially and temporally (Gaggiotti et al., 2018;
González-Megías et al., 2007). However, the
factors that drive biodiversity variation among
different groups of organisms are not fully
understood (González-Megías et al., 2007). Cli-
mate and anthropogenic activities, such as land
use changes, influence biodiversity (Aguirre-
Gutiérrez et al., 2017; Devictor et al., 2012; Fine,
2015). Understanding these effects is vital for
better biodiversity management and protection.
Butterflies are among the most diverse
insect orders globally (Kawahara & Breinholt,
2014). Despite this, conservation efforts for
butterflies, particularly in the tropics, are lim-
ited. Increasing knowledge about their popu-
lations and the factors influencing them is
crucial (Dunn, 2005; Sánchez-Bayo & Wyck-
huys, 2019). Moths and butterflies rely heavily
on plants and are sensitive to environmental
factors like temperature, humidity, light, and
habitat structure, which affect their life cycles
and behaviors (Aguirre-Gutiérrez et al., 2017).
Their sensitivity makes them excellent bioin-
dicators. Butterflies’ presence, abundance, and
diversity reflect ecosystem health and quality
(Fleishman & Murphy, 2009; Oostermeijer &
Van Swaay, 1998). Additionally, butterflies are
easy to monitor due to their diurnal activity,
abundance, and well-documented taxonomy,
making them valuable for assessing biodiversity
and environmental changes.
At the local level, elevation, habitat, and
seasonality influence butterfly diversity (Agu-
irre-Gutiérrez et al., 2017; Camero & Calderón,
2007; Carrero et al., 2013; Monteagudo et al.,
2001; Palacios & Constantino, 2006; Stephen &
RESUMEN
Efecto de la elevación, el hábitat y la estación en los ensamblajes de mariposas
(Lepidoptera: Papilionoidea) en una montaña tropical
Introducción: La variación espacial y temporal en el ambiente promueve la biodiversidad biológica. Sin embar-
go, los efectos combinados de la elevación, la estacionalidad y el hábitat sobre la diversidad y abundancia de
mariposas aún no han sido completamente explorados en ecosistemas tropicales rurales. Las mariposas, como
bioindicadores, ofrecen una excelente oportunidad para evaluar los impactos ambientales debido a su sensibilidad
a los cambios en el hábitat.
Objetivo: Este estudio tuvo como objetivo evaluar la variación en la abundancia de mariposas y la riqueza
de especies a través de diferentes hábitats, elevaciones y estaciones en un paisaje montañoso tropical rural en
Costa Rica.
Métodos: El estudio se llevó a cabo durante seis meses, abarcando tanto la temporada seca como la lluviosa. Se
muestrearon mariposas a lo largo de un gradiente elevacional (1 200, 1 500 y 1 800 m) en dos tipos de hábitat: pra-
deras y bosques riparios. Los métodos de muestreo incluyeron redes manuales y cebos de frutas en cada transecto
y en cada sitio durante seis meses. Se realizaron seis visitas a cada sitio, con siete transectos muestreados por visita
para capturar mariposas. Se registraron un total de 1 421 individuos representando 151 especies.
Resultados: La familia Nymphalidae presentó la mayor riqueza de especies y abundancia. La abundancia de
mariposas disminuyó con el aumento de la elevación, pero los patrones de riqueza de especies variaron según
la familia. La variación estacional tuvo un efecto significativo sobre la abundancia y la riqueza, con valores más
altos registrados durante la temporada de lluvias. Según la abundancia local, las especies raras se encontraron
más comúnmente en bosques riparios a mayores elevaciones durante la temporada húmeda. El hábitat no influyó
significativamente en la abundancia general de mariposas ni en la riqueza.
Conclusiones: La estacionalidad y la elevación son factores clave que influyen en las poblaciones de mariposas
en los ecosistemas montañosos tropicales. La falta de un efecto significativo del hábitat sugiere que otras variables
ecológicas pueden mediar los impactos del hábitat. Se requieren más estudios para esclarecer estas dinámicas,
particularmente el papel de la heterogeneidad del hábitat.
Palabras clave: gradiente elevacional, ecología del paisaje, patrones de biodiversidad, ecología de poblaciones,
bosque tropical, Lepidoptera.
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Sánchez, 2014). Diversity tends to decrease at
higher elevations (Camero & Calderón, 2007;
Carrero et al., 2013; Monteagudo-Sabaté et al.,
2001). Landscape structure also impacts butter-
fly abundance and richness (Aguirre-Gutiérrez
et al., 2017). It remains unclear whether slightly
disturbed environments support greater diver-
sity than more heavily disturbed ones (Concha-
Bloomfield & Parra, 2006; Tobar & Ibrahim,
2010). Additionally, seasonality influences but-
terfly community structure, with higher diver-
sity observed during months of higher rainfall
(Braby, 1995; Grøtan et al., 2012; Spitzer et
al., 1993). The influence of these factors may
vary by site or taxonomic group, as different
families or species may respond differently to
local conditions. This variation highlights the
importance of studying these factors at family
and regional levels.
Studying butterfly population variation in
tropical areas, considering environmental fac-
tors like habitat fragmentation, climate change,
and seasonal fluctuations, can provide valuable
insights into their conservation (Bonebrake et
al., 2010; Bonebrake & Deutsch, 2012). The
aim of this study was to examine the variation
in butterfly richness and abundance across
seasons (dry-wet), elevational gradients (1 200
to 1 800 m above sea level), and habitat types
(grassland and riparian forest) in a mountain
landscape in Costa Rica.
MATERIALS AND METHODS
The study was carried out in San Miguel
de Grecia, Costa Rica (10°07’07.85” - N
84°17’30.24” W) (Fig. 1), in an elevational
gradient from 1 200 to 1 800 meters above sea
level, between the Vigía and El Monte rivers.
The temperature ranges between 18º and 28°C
(Solano & Villalobos, 1996); the rainy peri-
od occurs between May and November, and
the dry period occurs between December and
April (Municipalidad de Grecia, 2003; Instituto
Meteorológico Nacional [IMN], 2018). The
area is located in the lower montane humid
forest life zone (Holdridge, 1967) on the Pacific
side of the country. The site consists of exten-
sive secondary forests, fragments of mature
forests, crops (coffee, tomato, and chili), and
livestock, which is thus a typical rural area.
We selected two types of habitats: grass-
lands and riparian forests. The riparian forest is
characterized by vegetation on the river’s edge
Fig. 1. Map of the San Miguel, Grecia, Alajuela, Costa Rica sampling stations.
4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64533, mayo 2025 (Publicado May. 15, 2025)
consisting of oak trees (Quercus sp.), Heliconia
sp., and Melastomataceae. At 1 200 m, the veg-
etation is more deforested compared to similar
forests at other elevations. During the dry sea-
son, the river continues to flow, but it decreases
to 1 200 m and 1 800 m due to water extraction
for agricultural irrigation and livestock use. The
grasslands are dominated by grasses for cattle
and other plants such as Solanum myriacan-
thum (Solanaceae), scattered custard apple trees
(Annona cherimola), and fig trees (Ficus spp.).
The landscape structure of this site is affected
by seasonality since most of the herbs have
considerably reduced foliage in the dry season.
We conducted fieldwork between Febru-
ary and September 2018. We established six
sampling stations at three elevations: 1 200 m,
1 500 m, and 1 800 m. At each elevation, we
selected two sites representing qualitatively
distinct habitats: grasslands and riparian forest
(Fig. 1). At each site, we established three tran-
sects per habitat (grasslands and riparian for-
est), resulting in six transects per elevation and
18 transects in total. Each transect measured 75
m in length and 4 m in width.
We conducted sampling on three consecu-
tive days per month, with one day dedicated
to sampling the transects at each elevation.
Each sampling day started at 7:00 h and ended
at 13:00 h, with 15 minutes of active observa-
tion per transect. At each transect, two people
conducted qualitative observations and sys-
tematically collected data. The two individu-
als walked simultaneously along the transect,
maintaining a separation of 2 meters to cover
the 4-meter width. The observation time at
each transect was 15 minutes, during which we
recorded all relevant observations (e.g., species
sighted, behavior). After completing a transect,
we allocated 30 minutes to move to the follow-
ing sampling site within the same elevation. In
total, we sampled 18 hours per site (3 days of 6
hours each). Considering the six sites, the effec-
tive sampling time accumulated by the end of
the study was 108 hours.
We used a butterfly net 2 m long and 0.5 m
in diameter for capturing specimens. We placed
two posts per transect, one at 0 m and the other
at 75 m; we used a banana as an attractant
(Baker & Baker, 1975), which was hung on each
post a day before sampling. The bananas were
left to ripen for two weeks before being used
as bait. We trapped all butterflies encountered
during the 15-minute round trip on the tran-
sect and temporarily placed them in a cloth
bag. At the end of the sampling period, one
individual of each species was retained for later
identification in the Entomology Laboratory of
the School of Biology, University of Costa Rica,
using Lamas (2004), DeVries (1987), DeVries
(1997), and Warren et al. (2013). At least one
individual per species was vouchered in the
Museo de Zoología at the Universidad de Costa
Rica (MZCR).
We summarized the data by habitat, eleva-
tion, and season and estimated species richness
and abundance for each condition. To compare
the richness between elevations, habitats, and
seasons, we used rarefaction analysis using Hill
numbers of order 0, 1, 2 (Chao et al., 2014).
To determine the effect of season, habitat, and
elevation on the abundance and richness of but-
terfly species, both in the overall data and for
each family, we used generalized linear models
(GLM) with the Poisson distribution because
the frequencies are low counts.
For the analysis of abundance by family, we
selected only those families with more than 20
individuals and those with more than five spe-
cies for the richness analysis. The significance
of each variable (habitat, elevation, and season)
was tested in the models using maximum like-
lihood tests. We analyzed the similarity of the
structure of butterfly assemblages according to
elevation, habitat, or seasonality using a Bray-
Curtis similarity index. We plotted the distanc-
es using a non-metric multidimensional scaling
(NMDS) ordination method. We performed all
analyses using the R program (version 3.5.3) (R
Development Core Team, 2022).
RESULTS
We found 1 421 butterflies belonging to
six families, 20 subfamilies, 96 genera, and 151
species. Nymphalidae was the best-represented
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family, with 936 individuals in 85 species and
55 genera (Supplementary information: Table
1). In contrast, the other families found were
Pieridae, Hesperiidae, Lycaenidae, Papilionidae,
and Riodinidae; each family had fewer than 250
individuals and fewer than 30 species. The most
abundant species was Hermeuptychia hermes
(Satyrinae), with 119 individuals, followed by
Celastrina argiolus gozara (Lycaenidae), with
115 individuals. Only six species were recorded
in all habitats and during all months sampled:
Catasticta nimbice bryson, Dryas iulia, Heli-
conius clysonymus montanus, Heliconius erato
petiverana, Hermeuptychia hermes, and Morpho
helenor narcissus (Supplementary information:
Table 1). Most species (133 species) showed a
low abundance, between 1-10 individuals dur-
ing all six months of sampling, while 16 species
had abundances of more than 10 individuals
(Fig. 2). For 57 species, there was only a single
individual recorded during the entire sampling
period (Fig. 2).
Abundance: The number of individuals
found in each habitat was 706 for the riparian
forest and 698 for the grassland. Butterfly abun-
dance per transect was not significantly dif-
ferent between habitats (Table 1, Fig. 3A). We
Fig. 2. The frequency of individuals for each butterfly species ordered according to their abundance throughout the sampling
period. Some species’ names are labeled to identify some points.
Table 1
Generalized linear models of the abundance and richness
of butterflies according to habitat, altitudinal gradient, and
season.
Variable Residual
Desv. Df P (Chi)
Abundance H0591.10 35
Habitat 591.05 34 0.83
H0591.6 35
Season 389.6 34 < 0.001
Elevation 351.31 32 < 0.001
Richness H0313.39 35
Habitat 313.23 34 0.99
H0313.394 35
Season 112.313 34 < 0.001
Elevation 96.614 32 < 0.001
6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64533, mayo 2025 (Publicado May. 15, 2025)
recorded 121 individuals of Lycaenidae in the
riparian forest, mostly belonging to C. argiolus
gozara. We found more individuals of Hes-
periidae in the grassland than in the riparian
forest (Fig. 4A). The most abundant family in
both habitats was Nymphalidae, among which
Heliconiinae was more abundant in the ripar-
ian forest. In contrast, Satyrinae with H. hermes
Fig. 3. Distribution of the number of butterflies/transect according to habitat (A) and according to elevation and season (B).
Fig. 4. Abundance (A, B, C) and species richness (D, E, F) of butterflies by subfamilies according to habitat (A, D), season (B,
E), and elevation (C, F). Subfamilies are ordered according to abundance (A, B, C) or species richness (D, E, F).
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Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64533, mayo 2025 (Publicado May. 15, 2025)
Fig. 5. Abundance (A, B, C) and species richness (D, E, F) of butterflies by subfamily of Nymphalidae according to habitat (A,
D, season (B, E) and elevation (C, F). Subfamilies are ordered according to abundance (A, B, C) or species richness (D, E, F).
was the dominant species in the grasslands (Fig.
5A). In addition, a dominance of Nymphali-
nae individuals was found in the grasslands
(Fig. 5A).
On the other hand, within the elevational
gradient, we found the highest abundance (587
butterflies) at 1 200 m, followed by 433 individ-
uals at 1 800 m, and 401 individuals at 1 500 m
(Fig. 3B). Butterfly abundance per transect
differed significantly between elevations (Table
1, Fig. 3B), with the lower elevation having the
highest butterfly abundance. The family Lycae-
nidae was most abundant at 1 800 m, with 128
individuals, compared to the other two eleva-
tions, while we did not observe the families
Papilionidae and Riodinidae at 1 500 m (Fig.
4C). In general, Nymphalidae was the most
abundant family at all three elevations (Fig.
4C). Nymphalinae and Heliconiinae were most
abundant at 1 200 m, Satyrinae at 1 500 m, and
Danainae at 1 800 m. (Fig. 5C).
The abundance of butterflies was higher
during the rainy than the dry months (Fig. 4B),
with 972 individuals during the wet season and
449 during the dry season (Table 1, Fig. 3B).
During both seasons, Nymphalidae was the
family with a greater abundance. Lycaenidae
was more abundant during the dry season
(Fig. 4B). Conversely, within Nymphalidae,
the subfamilies Apaturinae and Limenitidinae
were absent during the dry season. (Fig. 5B).
Satyrinae was the only subfamily that remained
constant during the two seasons (Fig. 5B).
Heliconiinae was the most abundant subfam-
ily in the wet season, with 166 individuals, and
Dryas iulia was the most abundant species (Fig.
4B). In the wet season, we recorded 142 indi-
viduals of the Ithomiini subfamily, and Ithomia
heraldica was the most abundant species in this
subfamily (Fig. 5B).
Nymphalidae, Lycaenidae, and Pieridae
were the most abundant families (Fig. 4).
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However, only in Lycaenidae, habitat and ele-
vation affected abundance (Table. 2, Fig. 6).
Lycaenidae was significantly more abundant in
high elevations near the river. In contrast, the
abundance in the other sampling sites was very
low (Fig. 6). Elevation also affected the abun-
dance of Nymphalidae and Pieridae since we
found more individuals at 1 200 m. However,
habitat did not affect butterfly abundance in
any elevation (Table. 2, Fig. 6).
Species richness of butterflies: We found
110 species in the grassland compared to 113
species found in the riparian forest, indicat-
ing no significant effect of habitat on species
richness (Table 1). Nymphalidae was the most
diverse family in both habitats (Fig. 4D). In
both the riparian forest and the grasslands, the
second most diverse family was Pieridae (Fig.
4D). The subfamily Apaturinae was found only
in grassland, while Satyrinae had more species
in the riparian forest, although it was more
abundant in the grassland (Fig. 5D).
In the elevational gradient, we found 94
species at the lowest elevation. In contrast, at
1 500 m and 1 800 m, we found 77 and 81 spe-
cies, respectively, resulting in a significant effect
of elevation on species richness (Table 1). We
found higher species richness in Pieridae and
Nymphalidae at 1 200 m. Furthermore, we did
not find Riodinidae and Papilionidae species
at 1 500 m. Still, there were more Lycaenidae
species at 1 800 m (Fig. 4F). For all elevations,
the family with the highest number of species
was Nymphalidae (Fig. 4F). Within this family,
Danainae was primarily found at medium and
high elevations, while for Nymphalinae and
Heliconiinae, more species were found at 1 200
m (Fig. 5F).
In the rainy season, we found 142 species,
while 52 species were found in the dry season,
indicating that seasonality influenced the spe-
cies’ richness (Table 1). Pieridae and Nym-
phalidae showed greater abundance in the rainy
season (Fig. 6). Nymphalidae had the highest
number of species in both the rainy and dry
seasons. Still, we did not find Apaturinae and
Limenitidinae species during the dry season
(Fig. 5E). Danainae was the most diverse sub-
family during the rainy season (Fig. 5E).
According to the rarefaction analysis, in
terms of total species, no significant differ-
ence was found between the riparian forest
and grassland habitats (Fig. 7A). However, the
Tabl e 2
Generalized linear model of the abundance and richness by family butterflies according to season and elevation, for the most
abundant families.
Family Variable Residual Desv. Df P (Chi)
Abundance Nymphalidae H0556.01 35
Habitat 552.16 34 0.050
Elevation 491.97 32 < 0.001
Lycaenidae H0411.12 35
Habitat 287.97 34 < 0.001
Elevation 116.61 32 < 0.001
Pieridae H0232.04 35
Habitat 230.30 34 0.23
Elevation 218.12 32 < 0.001
Richness Nymphalidae H0352.21 47
Habitat 352.20 46 0.95
Elevation 345.60 44 < 0.05
Pieridae H0121.66 47
Habitat 121.20 46 0.49
Elevation 114.14 44 < 0.05
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grassland was richer in common and abundant
species than the river, where there were more
rare species (Fig. 7A). We observed 39 species
in the river absent from the grassland, and 40
species in the grassland absent from the river.
On the other hand, at 1800 m, we found rarer
and less abundant species compared to the dif-
ferent elevations (Fig. 7B). Also, the most abun-
dant and common species were found at 1200
m and 1500 m, with similar species richness
at both elevations. In addition, there were sig-
nificant differences in richness between the dry
and rainy seasons (Fig. 7C), and we observed
rarer species in the rainy season (Fig. 7C).
Regarding the species richness within the
most diverse families, Nymphalidae had more
species at 1 200 m than 1 800 m. However, spe-
cies richness did not differ between habitats.
Pieridae had more species at 1 200 m compared
to the other elevations, with habitat having no
significant effect on species richness (Table
2). The other four families identified in the
sampling did not show differences in species
richness with respect to elevation and habitat.
Seasonality was the variable with a more
significant effect on the species composition of
the butterfly assemblages (r2 = 0.51, p = 0.003).
At the same time, we did not find an effect of
habitat (r2 = 0.07, p = 0.51) or elevation (r2 =
0.25, p = 0.26) on the species composition of
the butterfly assemblages (Fig. 8). The results
were similar if we considered the abundance
of the species (habitat: r2 = 0.13, p = 0.24; sea-
son: r2 = 0.52, p = 0.003; elevation: r2 = 0.18,
p = 0.43).
DISCUSSION
Compared to other larger sites in Costa
Rica with similar habitats, we observed a
high diversity of butterflies (Córdoba-Alfa-
ro, 2011; DeVries, 1991; Tobar et al., 2006;
Vega, 2012). Nymphalidae is one of the most
diverse butterfly families (Chacón & Montero,
Fig. 6. Distribution of the abundance of butterflies/transect/day of the Nymphalidae, Lycaenidae and Pieridae, according to
elevation and habitat.
10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64533, mayo 2025 (Publicado May. 15, 2025)
2007), which explains its prominence in this
study, while Riodinidae and Papilionidae were
observed with lower abundance. As Owen et
al. (1972) mentioned, these families might have
low abundance. However, more species could
emerge with increased sampling efforts, as
these two families have been previously record-
ed at this site (Murillo-Hiller, 2018).
Fig. 7. Rarefaction curves of butterfly species according to habitat (A), elevational gradient (B), and season (C).
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Among the most abundant species, H.
hermes stands out for using grasses (Poaceae)
as a host plant (Janzen & Hallwachs, 2009), a
major component of grasslands. We observed
over 100 C. argiolus gozara (Lycaenidae) indi-
viduals in the riparian habitat, primarily during
the dry season. This species has been reported
as highly seasonal in Oregon, USA (Warren,
2005). We also observed male aggregations on
mud and excrement and downstream migra-
tions at the end of the dry season. Butter-
fly communities in tropical forests typically
have high species richness but low abundance
(Owen, 1971). In our study, 35.6% of the total
sample consisted of species, with only one indi-
vidual recorded throughout the study period.
This pattern aligns with Pozo et al. (2008), who
state that rare species determine monthly varia-
tion in species richness. Among the rare species
observed, Forsterinaria neonympha, Dynamine
paulina thalassina, and Pteronymia hara are
rarely recorded in Costa Rican collections,
highlighting the importance of studying new
sites to detect range expansions.
No significant differences in butterfly
diversity were found between habitats. The
grassland contained features such as living
fences and a rich diversity of herbaceous plants,
which enhanced connectivity between forest
fragments (Enrique-Tobar & Ibrahim, 2010;
Ospina-López et al., 2015). This vegetation
diversity provides reproduction and feeding
sites for species more typical of riparian forests,
such as Mechanitis menapis saturata (Ithomi-
ini), which laid eggs on Solanum myricanthum
(Solanaceae) in the grasslands.
The riparian forest is a highly hetero-
geneous habitat where the river’s dynamics
modify vegetation, causing frequent light and
temperature changes. These modifications may
lead to greater specialization among species
dependent on riparian flora, but they also sup-
port generalist species, contributing to higher
butterfly diversity (Naiman & Decamps, 1997;
Naiman et al., 1993). Species found only in
the riparian forest include C. argiolus gozara,
Apuelca maeonis, Atlides sp. (Lycaenidae), and
Manataria hercyna maculata, among others.
Fig. 8. Ordination of the butterflies’ assemblages according to habitat, season and elevation using Bray-Curtis similitude
indexes and non-metric multidimensional scaling.
12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64533, mayo 2025 (Publicado May. 15, 2025)
Although no significant differences in
richness and abundance were found, the exclu-
sivity of species to each habitat suggests that
landscape heterogeneity is crucial for main-
taining biodiversity. Grasslands and riparian
forests, with their unique species richness and
abundance, contribute to increased taxonomic
and functional diversity in the ecosystem. Dis-
turbances, particularly in fragile habitats like
riparian forests, can affect populations depen-
dent on these areas.
Various abiotic factors influence species
diversity along elevational gradients in the
tropics and are complex (Monteagudo-Sabaté
et al., 2001). The general trend is that species
richness decreases with increasing elevation,
though different taxonomic groups may exhibit
independent variations (Andrade, 1998). We
observed that Lycaenidae was more abundant
at 1 800 m, where forest cover was higher, while
Pieridae and Nymphalidae showed greater rich-
ness and abundance at 1 200 m.
We found that seasonality affects the abun-
dance, and the diversity observed, with higher
butterfly abundance during the rainy season
and the highest species number recorded in
July. This pattern has been observed in other
neotropical regions (Checa, 2006; Checa et al.,
2009; Pozo et al., 2008) and on the Pacific slope
of Costa Rica (Janzen, 1993; Murillo-Hiller et
al., 2019). During the dry season, many but-
terflies remain in the pupal stage, which is more
drought-resistant (Janzen, 1993). The first rain
triggers adult emergence, coinciding with
increased foliage and resources for reproduc-
tion (Barth 1991; Frankie et al., 1976; Proctor
et al., 1996; Williams-Linera & Meave, 2002).
However, phenological patterns are usually
complex and cannot be generalized since some
species can show a greater abundance during
the dry season or are unaffected by seasonality
(Gilbert & Singer, 1975). Resource availability
can vary for each species throughout the year,
and some herbs produce more resources for
butterflies during the rainy season (Fenner,
1998). This heterogeneity promotes the diver-
sity and abundance of butterflies found at the
sample sites throughout the year.
Climatological anomalies could influ-
ence butterfly populations (Srygley et al.,
2010; Grøtan et al., 2014). Grøtan et al. (2014)
observed that butterfly diversity had peaks in
biannual cycles, affected by the abundance of
plant resources, which in turn was affected
by precipitation. During the present study,
the phenomenon of La Niña influenced the
weather conditions during the sampling period
(Zhang et al., 2019). Among the effects that this
phenomenon exerts is an increase in rainfall;
consequently, the butterfly’s assemblage could
vary in their phenology (Wallis DeVries et al.,
2011; Wilson & Maclean, 2011).
This study highlights the importance of
seasonality and elevation in monitoring popu-
lations, as these factors influence species dis-
tribution and conservation. We found higher
butterfly abundance during the wet season
and at lower elevations, with riparian forests
and grasslands showing similar abundance.
These findings underscore the critical need for
effective conservation strategies that consider
seasonal and elevational factors and the preser-
vation of habitat diversity. Protecting areas such
as riparian forests and grasslands is vital for
maintaining butterfly populations and ensuring
the resilience of these ecosystems in the face of
climate change and habitat disturbances. Con-
servation efforts should focus on preserving
landscape heterogeneity to support biodiversity
and prevent the loss of species that depend on
these unique habitats.
Ethical statement: the authors declare that
they all agree with this publication and made
significant contributions; that there is no con-
flict 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 Germán Vega from the National
Museum of Costa Rica and Bill Haber for their
13
Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64533, mayo 2025 (Publicado May. 15, 2025)
help in identifying butterflies. Thanks also to
the landowners where we did the fieldwork. We
also thank Jeiner Matamoros, Mayela Calde-
rón, and Liseth Matamoros for their financial
collaboration towards this project. We thank
Natalia Jiménez Conejo for her contributions
to our study. Finally, we thank two anonymous
reviewers who improved the paper and the text.
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