Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64530, mayo 2025 (Publicado May. 15, 2025)

Diversity of moths and butterflies of the southwestern region of Costa Rica

Isidro Chacón1; https://orcid.org/0000-0001-5139-2073

Ana Catalina Sánchez-Quirós2; https://orcid.org/0000-0002-8714-4327

Gilbert Barrantes2, 3*; https://orcid.org/0000-0001-8402-1930

1. BioAlfa, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, Costa Rica;

boconera@gmail.com

2. Escuela de Biología, Universidad de Costa Rica, San José, Costa Rica; catasq@gmail.com

3. Centro de Investigaciones en Biodiversidad y Ecología Tropical, Universidad de Costa Rica, San José, Costa Rica.

gilbert.barrantes@gmail.com (*Correspondence)

Received 01-IX-2024. Corrected 30-XII-2024. Accepted 04-III-2025.

ABSTRACT

Introduction: The Costa Rican southwestern region is considered one of the neotropical biodiversity hotspots.

This geographically isolated forest embraces a rich diversity of moths and butterflies, which is still unknown for

the most part.

Objectives: (a) Describe and compare the diversity of moths and butterflies among different localities of the

southwestern Costa Rican region, and (b) identify information gaps in this group of insects.

Methods: Data for this study were collected in 12 different localities by parataxonomists and taxonomists for

the national inventory of biodiversity led by INBio (National Biodiversity Institute) using different trap types,

but relying mainly on light traps. We compared alpha and beta diversity of moths and butterflies among the six

localities (Cortés, Piedras Blancas, Rancho Quemado, Agujas, Los Patos, Sirena) with more extensive sampling.

Results: The dataset consisted of 78 747 specimens, of which 2 096 were identified only to the order level. The

remaining 76 650 specimens were classified to the family level (n = 48 families), genus, species, or morphospe-

cies. Species and morphospecies were distributed across 37 families. The alpha and beta diversity varied across

localities for the 12 families of moths and butterflies that were present in the six localities selected. In general,

Piedras Blancas, Rancho Quemado, and Sirena stand out as the most diverse localities. The richness of species

varied among different families. For most families Agujas, Los Patos, and Sirena contained more species, but

some other families showed a different pattern.

Conclusion: Differences in the diversity of moths and butterflies over a northern-southern gradient likely cor-

respond to species (individuals within species) adapted to different biotic (e.g., availability of host plants and

food resources) and abiotic (e.g., microclimatic conditions associated to the complex topography of the region)

conditions. Despite the large effort made by INBio in knowing and mapping the biodiversity of Costa Rica, and

the economic and biological importance of the biodiversity, there are still a huge number of species to be known,

named, and properly used.

Keywords: alpha diversity; beta diversity; moth and butterfly distribution; dissimilitude.

RESUMEN

Diversidad de polillas y mariposas en la región suroeste de Costa Rica

Introducción: La región suroeste de Costa Rica es considerada una de las áreas más diversas en el neotrópico.

Este bosque, geográficamente aislado, alberga una rica diversidad de polillas y mariposas, aunque en gran parte

aún desconocida.

https://doi.org/10.15517/rev.biol.trop..v73iS2.64530

SUPPLEMENT

SECTION: ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64530, mayo 2025 (Publicado May. 15, 2025)

INTRODUCTION

“Biological diversity must be treated more

seriously as a global resource, to be indexed,

used, and above all, preserved” (Wilson, 1988;

p. 3). These words so wisely and warningly

expressed by E. O. Wilson nearly four decades

ago, resound louder at the present time than

ever before. We continue losing species at an

unprecedented and accelerated rate, particu-

larly in the tropics (Gao et al., 2020; Janzen &

Hallwachs, 2021; Pievani, 2014). The factors

causing biodiversity loss are numerous and

most of them (if not all) have a destructive syn-

ergetic effect on ecosystem diversity. The factors

responsible for biodiversity loss can be visual-

ized as a complex network, in which habitat

destruction, fragmentation, urban expansion,

species invasion, contaminants (e.g., pesticides,

macro and microplastic, noise, light, etc.), and

climate change interact and affect differently

the community composition at each particular

time and ecosystem (Aguirre-Gutiérrez et al.,

2017; Sanchez-Azofeifa, 2000; Seymoure, 2018).

The destruction and fragmentation of large

extensions of tropical forests and other ecosys-

tems for agricultural and urbanization purposes

are certainly the primary factors responsible for

biological diversity loss (Joyce, 2006; Sanchez-

Azofeifa, 2000). Once natural ecosystems are

reduced to tracts of different size and with dif-

ferent degrees of deterioration, and separated

by different distances and barriers, other factors

that often have little impact in extensive natural

ecosystems (e.g., hunting, fishing, invasive spe-

cies, contamination) become a serious concern

for the preservation of the biodiversity in the

deteriorated tracts of tropical ecosystems (Cul-

len et al., 2000; Seymoure, 2018). The negative

impact of these factors is exacerbated by the

fluctuation and changes in the climatic condi-

tions caused by the climate change (Karmalkar

et al., 2008; Lyra et al., 2017).

In current discussions on biodiversity,

tropical forests continue occupying a central

stage, and a serious concern for its preservation

is the enormous number of species remaining

to be named (Erwin, 1982; Raven, 1988). We

are losing most species inhabiting this planet

without ever knowing them. These species are

not only important for the scientific commu-

nity, but they represent valuable resources for

the entire society (Burton et al., 1992; Smith et

al., 1992). The overwhelming task of catalog-

ing species is far behind, and two main factors

Objetivo: (a) Describir y comparar la diversidad de polillas y mariposas entre diferentes localidades de la región

suroeste de Costa Rica, e (b) identificar vacíos de información en este grupo de insectos.

Métodos: Los datos para este estudio fueron recolectados en 12 localidades diferentes por parataxónomos y taxó-

nomos para el inventario nacional de biodiversidad dirigido por el INBio (Instituto Nacional de Biodiversidad),

utilizando diferentes tipos de trampas. Se comparó la diversidad alfa y beta de polillas y mariposas entre las seis

localidades (Cortés, Piedras Blancas, Rancho Quemado, Agujas, Los Patos, Sirena) con un muestreo más extenso.

Resultados: El conjunto de datos consistió en 78 747 especímenes de los cuales 2 096 se identificaron solo a nivel

de orden. Los otros 76 650 especímenes se asignaron a familia (n = 48 familias), género, especie o morfoespecie.

Especies y morfoespecies fueron asignados a 37 familias diferentes. La diversidad alfa y beta varió entre localida-

des para las 12 familias de polillas y mariposas presentes en las seis localidades seleccionadas. En general, Piedras

Blancas, Rancho Quemado y Sirena destacaron como las localidades más diversas. La riqueza de especies varió

entre diferentes familias. Para la mayoría de las familias, Agujas, Los Patos y Sirena tuvieron más especies; algunas

otras familias mostraron un patrón diferente.

Conclusión: Las diferencias en la diversidad de polillas y mariposas a lo largo de un gradiente norte-sur proba-

blemente responden a especies (individuos dentro de las especies) adaptadas a diferentes condiciones bióticas (ej.:

disponibilidad de plantas huésped y recursos alimentarios) y abióticas (ej.: condiciones microclimáticas asociadas

a la compleja topografía de la región). A pesar del gran esfuerzo realizado por el INBio para conocer y mapear la

biodiversidad de Costa Rica, y la importancia económica y biológica de la biodiversidad, aún hay un gran número

de especies por conocer, nombrar y utilizar adecuadamente.

Palabras clave: diversidad alfa; diversidad beta; distribución de polillas y mariposas; disimilitud.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64530, mayo 2025 (Publicado May. 15, 2025)

worsen this enormous task. First, worldwide

governmental institutions cut (or eliminate)

funding resources for taxonomy and natural

history research, second and directly linked to

the previous factor, taxonomists are rarer and

rarer, in part because taxonomy and systematics

are, at present, considered irrelevant study areas

within the broad scope of biology. The scien-

tific society, and the society in general, need to

pause and rethink the importance of naming

and preserving the species on the planet. Sci-

ence has found and continues finding new uses

for biological diversity, but without knowing a

simple but important thing –the species name–

we may be losing important resources. “We

must know that [species] exist before we can

understand or use them” (Raven, 1988, p. 120).

We show the problem that concerns bio-

diversity preservation by focusing on diver-

sity of moths and butterflies in one the most

diverse areas in the neotropics, the central and

southwestern Costa Rican region (southwest-

ern region hereafter) including the Península

de Osa, a diversity hotspot (Cornejo et al., 2012;

Haffer, 1974; Morera-Beita et al., 2019). This

region embraces one of the most diverse low-

land neotropical forest, with a large number

of endemic species (Gilbert et al., 2016). It has

been the focus of a large number of investiga-

tions in different groups of organisms, but it

has also been subject to intense destruction

and fragmentation as a consequence of timber

exploitation (usually illegal), introduction of

different crops (often unsuccessful), and gold

mining (Gilbert et al., 2016; Lobo et al., 2007),

which have gradually eroded the rich biodiver-

sity of the region. In an attempt to show the

diversity that fragments of this magnificent

forest still maintain, we studied the diversity

of moths and butterflies in six localities dis-

tributed over the southwestern region of Costa

Rica. Lepidoptera is the third or fourth most

diverse order of insects with over 155 000

described species (Eggleton, 2020), and they

play diverse and important roles in the ecosys-

tems (Eggleton, 2020; Ghazanfar et al., 2016).

Larvae of nearly all moths and butterflies are

herbivorous, with many highly specialized on

one or a few plant species, and as adults they

often serve as pollinators of different groups

of plants, including some crops (Halder et al.,

2019). From an evolutionary perspective, moth

and butterfly herbivory and pollination are evo-

lutionary forces that have shaped the evolution

of entire groups of plants, and from an eco-

logical perspective moths and butterflies inter-

act with diverse groups (e.g., plants, parasites,

predators), becoming a key component for the

functioning (e.g., energy flow) of the ecosys-

tem. Therefore, losing moth-butterfly diversity,

entails not only species disappearances, but also

the disappearance of the functions and interac-

tions these insects play in the ecosystem.

In this study we focus on the following two

objectives. (a) To describe and compare the

diversity of moths and butterflies among differ-

ent localities of the southwestern Costa Rican

region, and (b) to identify the information gaps

in this group of insects. For the second objec-

tive we specifically focused on the number of

specimens which could not be assigned to a

particular species, which could be primarily

attributed to the fact that taxonomists are “a

species in extinction”, and to the limited funding

for taxonomy and systematics. To address these

objectives, we used a data base from a monitor-

ing project conducted in the region by INBio

(National Institute of Biodiversity). This is the

most comprehensive data base of moths and

butterflies for a lowland neotropical rainforest.

MATERIAL AND METHODS

Study sites: The study sites are in the

southwestern region of Costa Rica, and include

different forest types: wet forests, tropical moist

forests, and tropical premontane wet forests

(Holdridge, 1967), along an elevation from sea

level to 745 m on Cerro Rincón. The geomor-

phology of the region is complex with narrow

ridges, pronounced slopes, and sediment plains

(Gilbert et al., 2016; Weissenhofer & Huber,

2001). The annual precipitation ranges from

4 000–6 000 mm, and the mean annual temper-

ature fluctuates between 25 and 27 °C (Gilbert

et al., 2016). The rainy season occurs between

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64530, mayo 2025 (Publicado May. 15, 2025)

May and December, with a notable reduction

in precipitation between January through April.

Geomorphological and climatic conditions

have been determinant factors influencing the

complexity of the forest mosaic in the region.

Moths and butterflies were collected using

different procedures (see below) in 12 different

localities from 1973 to 2013, at the beginning

only a few localities were sampled sporadically.

However, from 1978 to 2013 samplings were

more frequent and intense and included more

localities. Each locality was sampled at irregu-

lar time intervals during the sampling period.

Selection of localities was based on two criteria:

that they were in a protected area (e.g., national

park, biological reserve), and with access to

electricity or an electric generator. Electricity

was necessary to charge batteries to set light

traps and prepare moth and butterfly speci-

mens for later identification. All localities were

within large tracts of protected mature lowland

rainforest. This survey was part of the national

biodiversity inventory led by INBio (National

Biodiversity Institute). Field collections were

carried out by parataxonomists and taxono-

mists, and identification of specimens was done

by local and foreign taxonomists; names and

affiliation of specialist taxonomists for each

family are included in Supplementary Table 1.

All specimens were catalogued and deposited

at the INBio entomological collection, currently

deposited at the Museo Nacional de Costa Rica.

Collecting procedures: Parataxonomists

used three different methods to collect butter-

flies: light traps, fruit traps, and entomological

net. Light trap was the method more exten-

sively used and consisted of setting the traps

for 5 nights/month (two before new moon,

new moon, and two after new moon); these

collecting methods are explained in detail by

Janzen (1983). Samplings were non-systematic

and were conducted at different time periods in

each locality. In each locality, parataxonomists

set 1 or 2 light traps, and 15 sampling stations

for fruit traps. Each fruit trap station consisted

of 2 traps, one set in the understory (at 2 m

from the ground) and the other in the canopy

(~ 20 m high). Despite the inconsistent sam-

pling, such valuable data are still comparable

using the appropriate statistical procedures (see

Statistical analyses section). We excluded speci-

mens captured with Malaise traps, because this

method was used only in one site.

Statistical analyses: Richness surveys

(INBio biodiversity inventory) are usually

focused on collecting the most species possible

in a particular locality; thus, different sam-

plings in the same locality often vary spatially

and temporarily (moving from one location

to another in the same site, or during different

times of the year). This is in contrast to system-

atic sampling procedures in which samplings

are carried out in the same locations with a

specific duration of time, but systematic sam-

plings limit the number of species detected in

a particular site. Hence, given the conditions of

this inventory, we compared diversity param-

eters across sites, rather than between periods

within sites.

We selected six different localities as the

focal sampling units (Fig. 1; Table 1). Selection

of these sites was based on coverage (sampling

completeness – see Alpha diversity section), but

data from other sites (indicated when neces-

sary) were included in more general analyses

(e.g., total number of species).

Beta Diversity: For beta diversity anal-

yses, we selected 12 families (Apatelodidae,

Bombycidae, Crambidae, Erebidae, Geometri-

dae, Lasiocampidae, Megalopygidae, Noctui-

dae, Notodontidae, Nymphalidae, Saturniidae)

that were all present in the six localities most

extensively sampled (Cortés, Piedras Blancas,

Rancho Quemado, Agujas, Los Patos, Sirena;

Fig. 1). We provided general comparative infor-

mation on the number of species in the families

selected. We also compared the dissimilarity of

species for each family among sites using the

Sorensen index. We conducted this analysis in

two steps: first, we estimated the dissimilarity

of species within each family among localities;

second, we calculated the mean and 95%-con-

fident intervals for each family, and compared

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64530, mayo 2025 (Publicado May. 15, 2025)

them across the 12 families. Dissimilarity was

calculated with the function beta.pair from the

betapart package (Baselga, 2017), on a pres-

ence/absence data set. This type of data and

analysis notably reduce the effect of differences

of sampling effort among sites.

Alpha diversity: We calculated and com-

pared alpha diversity for all species (not only

for species in the six families included in

previous analyses) in each of the six focal

localities using the iNEXT () function, and

estimated the cover-based Hill diversity using

the estimateD () function of the iNEXT pack-

age (Chao et al., 2014; Hsieh et al., 2016). We

calculated three diversity metrics (q = 0, q

= 1, and q = 2): q = 0 estimates the species

richness and is more sensitive to sample size

and influenced by rare species; q = 1 provides

equal weight to rare and common species; and

q = 2 provides greater weight to the dominant

species (Chao et al., 2014; Hsieh et al., 2016).

Fig. 1. Map of the study region showing the distribution of the six localities (in yellow) used in the analyses: 1- Cortés, 2-

Piedras Blancas, 3- Rancho Quemado, 4- Agujas, 5- Los Patos, 6- Sirena.

Table 1

Geographical location of the six localities most extensively sampled during the study period in the southwestern region of

Costa Rica. The number of species and specimens collected in each of the six localities is included. Species and specimens

correspond to the 12 families selected (see Beta Diversity section in methods).

Locality Latitude Longitude No. species No. specimens

Cortes 9°, 58’ 83°, 31’ 469 1350

Piedras Blancas 8°, 42’ 83°, 16’ 1865 12870

Rancho Quemado 8°, 40’ 83°, 33’ 619 2356

Agujas 8°, 32’ 83°, 25’ 1837 13221

Los patos 8°, 31’ 83°, 34’ 654 1868

Sirena 8°, 28’ 83°, 35’ 2027 22575

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We used confidence intervals to evaluate the

differences across localities for each diversity

indicator (Chao et al., 2014). We also generated

rarefaction and extrapolation curves (ggiNEXT

function, iNEXT package) for each site, with

a 95% confidence interval (Chao et al., 2014).

The curves were estimated based on the abun-

dance method, for both coverage and sampling

units, to determine the completeness of col-

lections in each site. We then standardized

the samples by coverage to reduce the effect

of uneven sampling effort between localities,

allowing diversity measures to be more compa-

rable among sites.

Information gaps: We provided compara-

tive tables to account for the information gaps

of the study group. We used the R statistical

language, version 4.4.0 (R Core Team, 2024) for

all statistical analyses.

RESULTS

The dataset consisted of 78 747 catalogued

specimens. Of these, 2 097 were identified only

to the order level. The remaining 76 650 speci-

mens were classified to the family level (n =

48 families), genus, species, or morphospecies.

Species and morphospecies were distributed

across 37 families (Supplementary Tables 2 and

3). Specialist taxonomists for each family iden-

tify the collected specimens, ensuring that all

specimens assigned to a specific morphospe-

cies belong to recognized taxonomic species

or, in many cases, to new, undescribed species

(Supplementary Table 1). Moths and butterflies

were collected at 12 different localities with a

varying capture effort in each one, but to make

data comparable, we focused on six localities

(Cortés, Piedras Blancas, Rancho Quemado,

Agujas, Los Patos, Sirena).

Beta diversity: The number of species var-

ied considerably among the six localities (X2 =

2116.1, df = 5, P < 0.001). This result was likely

influenced by differences in sampling effort,

since the number of species collected in each

locality correlated with the number of sampling

days (r2 = 0.86, P = 0.028). In these six localities

the two most species-rich families were consis-

tently Erebidae and Geometridae, although the

number of species of both families was greater

in Agujas, Los Patos and Sirena (Fig. 2a). Simi-

larly, Noctuidae and Notodontidae had more

species in these three sites. But richness of other

families did not have an apparent pattern with

respect to the number of species across locali-

ties. The proportion of species by family also

differed significantly (X2 = 535.33, df = 55, P

< 0.001; Fig. 2a) across localities and this test

is much less influenced by uneven sampling,

indicating that some local factors are likely

affecting differently the species richness in dif-

ferent families.

The dissimilarity (values vary between 0

- total similarity to 1- total dissimilarity) com-

paring species within families across locali-

ties, was, in general, low for all families (Fig.

2b). The maximum mean dissimilarity (0.33)

was obtained for Noctuidae, indicating that in

average about 70% of Noctuidae species were

shared among communities. Apatelodidae and

Bombycidae, followed by Saturniidae presented

a high similarity (low dissimilarity) of species

among localities. Particularly Apatelodidae and

Bombycidae had relatively few species, but they

were present in all (or nearly all) localities, sug-

gesting that these species have a wide distribu-

tion and they are adapted to a wide range of

conditions (e.g., habitats, diet).

Noctuidae, Erebidae, Lasiocampidae, and

Nymphalidae were species-rich families, which

shared fewer species across localities (Fig. 2b).

Species-rich families are usually composed of

species that range from extremely rare to very

common, and from specialists to generalists.

Such conditions restrict some species to a par-

ticular microhabitat that may not be present

in all localities or making species so rare that

it drastically reduced the capture probabil-

ity. Comparisons between all family pairs are

included in Supplementary Table 4.

Alpha diversity: For this analysis we

compared alpha diversity based on coverage,

including all species in each of the six localities

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64530, mayo 2025 (Publicado May. 15, 2025)

(Cortés, Piedras Blancas, Rancho Quemado,

Agujas, Los Patos, Sirena). Localities differed

for the three diversity indices (Richness, Hill-

Shannon, and Hill-Simpson). The coverage

(sampling completeness) was high and similar

for the six localities (Fig. 3a), allowing for a fair

comparison among them.

Species richness (q = 0) differed among

localities, with Piedras Blancas, Rancho Que-

mado, and Sirena standing out from the other

Fig. 2. Comparisons of number of species of moths and butterflies. a- Number of species of 12 families in six localities. b-

Comparison of dissimilitude among 12 families. For each family we obtained the mean and the 95% confident interval of

the dissimilitude of species in each family present in the six localities (Cortés, Piedras Blancas, Rancho Quemado, Agujas,

Los Patos, Sirena).

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localities (Fig. 3). These three localities had

the largest number of species (estimated by

coverage), but also included a wide range of

rare and common species, particularly Piedras

Blancas and Rancho Quemado; high estimates

of Hill-Shannon indicate a high number of spe-

cies, both rare and common. The Hill-Simpson

index (q=2), which is influenced by abundant

species, was higher for Rancho Quemado and

Piedras Blancas (Fig. 3). This indicates that

these two localities had a large number of com-

mon species. But, Rancho Quemado, Piedras

Blancas, and Sirena were the localities with

more species that range from rare to common

(Hill-Shannon q = 1).

DISCUSSION

We found that the Costa Rican southwest-

ern lowland forests stand out as having one

of the most diverse moth and butterfly fauna

in the neotropical region, with more than

3030 species. The diversity (richness and abun-

dance) of moths and butterflies changes across

Fig. 3. Diversity of moths and butterflies in the southwestern region of Costa Rica. a- Diversity estimates separated by panels:

Richness (q = 0), Hill-Shannon (q = 1), and Hill-Simpson (q-2). b- Hill-diversity estimates among the six localities studied

in the southwestern region of Costa Rica. Error bars correspond to 95% CI.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64530, mayo 2025 (Publicado May. 15, 2025)

localities over a northern-southern gradient.

Community composition – species richness

and abundance– is not “static”, since it changes

temporally and spatially (Enkhtur et al., 2021;

Heimonen et al., 2013). Differences in commu-

nities over a geographical gradient or geograph-

ical mosaic can be attributed to historical and/

or ecological factors (Barrantes, 2009). Histori-

cal factors, such as past geology (Bagley & John-

son, 2014) and climatic events (including recent

events), have shaped the species composition

in most if not all communities (Haffer, 1974).

Ecological factors, through species interactions

(e.g., parasitism, predation, competition) or by

adaptation (of individuals of populations) to

different microhabitats are more important in

population dynamics, and thus in regulating

species abundances (Stiling, 1988; Templeton,

1989). These factors underlie richness and

abundance patterns of all communities.

Changes in richness and abundance

occurred over the area, but not all families

respond similarly. The region is an “island of

lowland rain forest” surrounded by dry forests

and the highest cordillera of the region, with

a dynamic history of geological and climatic

changes that have resulted in a mosaic of veg-

etation types (Gómez, 1986; Hofhansl et al.,

2019). Consequently, this complex vegetation

landscape has offered the opportunity for spe-

ciation (a region with high endemism) and

variation in community composition of differ-

ent organisms, through adaptation processes,

within a small geographical scale (e.g., butter-

flies- Gilbert, 1973 plants- Hofhansl et al., 2019;

birds- Pereira & Barrantes, 2009).

Moths and butterflies, as diverse as this

group is, present an equally large range of

ecological requirements that range from

monophagous to polyphagous larvae (Brues,

1924; Chacón & Montero, 2007; DeVries, 1987;

Ehrlich & Raven, 1964). Similarly, the feeding

habits and habitat preference of adult moths

and butterflies range from highly specialized

(e.g., many Sphingidae species) to opportunis-

tic (generalist) species (Haber & Frankie, 1989).

Therefore, distribution of larval host plants,

flowering and fruiting plants used as food

resources by adult moths and butterflies, as well

as floristic diversity and forest structure (e.g.,

vegetation strata; DeVries, 1988), are key fac-

tors determining the richness, abundance, and

thus the distribution of butterflies and moths in

this study region.

Although detailed studies of the distri-

bution and abundance of host plants, and

resources distribution for adult moths and but-

terflies are lacking, it is possible to cautiously

make some extrapolation from small scale stud-

ies. Forest plant composition and vegetation

structure change over the region, determined

by soil, climatic, and topographic conditions

(Harms et al., 2001; Hofhansl et al., 2019). Birds

respond to topographic and vegetation varia-

tion in the region (Pereira & Barrantes, 2009).

Some species have a discontinuous distribution

(Corapipo altera, Ixothraupis guttata, Habia

atrimaxillaris) associated with some particu-

lar habitats or microhabitats. Distribution of

butterflies is tightly associated with vegeta-

tion structure which, in turn, correlates with

the topography (Binz et al., 2014). The lat-

ter authors found that nymphalids were more

diverse in ridge forests than in slopes and creek

forests. This is possibly associated with the

higher floristic diversity, which likely includes a

larger variety of host plants for larvae and food

resources for adults in ridge forests (Murdoch

et al., 1972, Novotny et al., 2007).

The variation in beta diversity shown in

this study may be associated with variation

in climate, topography, and vegetation along

the northern-southern gradient of localities.

Each locality may likely offer a different set of

resources for butterflies and moths, which may

further explain the different patterns of diver-

sity we detected across families. The local diver-

sity (for the three estimators of alpha diversity)

was higher in Piedras Blancas, Rancho Que-

mado and Sirena. These three localities not

only have the highest richness (Fig. 3A-B, but

also a large number of both rare and common

species (Fig. 3A-B). Although we used conser-

vative analyses (e.g., presence/absence data and

coverage instead of number of individuals) we

cannot completely reject the possibility that

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64530, mayo 2025 (Publicado May. 15, 2025)

our results are influenced by uneven sampling.

However, we are confident that the conservative

analyses we used show the general patterns of

diversity of moths and butterflies in the region.

For instance, the patterns of richness in some

families, such as Notodontidae and Nymphali-

dae, having a large number of species in locali-

ties with low sampling effort (Fig. 2), and the

proportion of species across localities, support

our argument.

Information gaps: Burton et al. (1992)

pointed out many “solid reasons” for the impor-

tance of preserving biodiversity: productivity,

pest control, aesthetic aspects, and more impor-

tantly – resources for the uncertain future

(e.g., undiscovered drugs and food sources).

More recently, the importance of insect pol-

lination has become an important issue for

maintaining crop production (Montero et al.,

2024). In Costa Rica, nature-based tourism is

attracted by high diversity places (Echeverri

et al., 2022). However, despite the economic

and social importance of preserving our diver-

sity, very few resources are assigned by private

(e.g., large hotels that directly benefit from the

biodiversity) and public institutions, for even

simple but important aspects, such as knowing

the Costa Rican biodiversity. The data used in

this study represent an important effort toward

this end.

In conclusion, the southwestern low-

land forested region of Costa Rica includes

an extremely rich diversity of moths and but-

terflies. This diversity varies among localities

and families. Piedras Blancas, Rancho Que-

mado and Sirena are the most diverse localities,

and Erebidae, Geometridae, Noctuidae and

Notodontidae stand out as the families with

most species. The patterns of diversity observed

are likely driven by variation of environmen-

tal conditions (e.g., topography, climate) that

influence vegetation structure and floristic

diversity, and so the variation and availability

of resources to which different species of moths

and butterflies (larvae and adults) are adapted

(Aguirre-Gutiérrez et al., 2017). Despite the

importance of knowing and protecting the

diversity in Costa Rica, very little effort has

been made to provide the resources for the

continuity of these studies (Camacho-Sandoval

& Duque 2001; Echeverri et al., 2022).

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.

ACKNOLEDGMENTS

We are grateful with all the group of

parataxonomists, taxonomists (local and for-

eign), and local persons that collected, identi-

fied, and catalogued all specimens of moths and

butterflies used in this study. We thank Karina

Montero, Nicole Gamboa, and Luis Sandoval

for advice in data analyses.

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