Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64528, mayo 2025 (Publicado May. 15, 2025)

Temporal changes in the diversity and abundance of stingless bee

nests in an urbanized environment

Jonas Konicek1; https://orcid.org/0000-0003-1464-5101

Wendolyn Matamoros-Calderón2; https://orcid.org/0000-0002-2715-1875

Mauricio Fernández Otárola2, 3*; https://orcid.org/0000-0001-9240-7569

1. Ulm University, Ulm, Germany. jonaskonicek@web.de

2. Biodiversity and Tropical Ecology Research Center (CIBET), University of Costa Rica, San José, Costa Rica.

wendolyn.matamoros@ucr.ac.cr

3. School of Biology, University of Costa Rica, San José, Costa Rica; mauricio.fernandez@ucr.ac.cr (*Correspondence)

Received 30-VIII-2024. Corrected 18-II-2025. Accepted 04-III-2025.

ABSTRACT

Introduction: Increasing urbanization has endangered many species to an unknown extent. Stingless bees

(Apidae: Meliponini) are highly important pollinators of tropical plants. Some species are well adapted to urban

areas and use man-made structures to build their nests. In Costa Rica, there are 59 stingless bee species, but no

account of their urban richness and abundance has ever been made.

Objective: To describe the composition and dynamics of the social bee community on the campus of the

University of Costa Rica in San José over a six-year period.

Methods: We systematically searched for stingless bee nests (active colonies) in trees, buildings, walls and other

man-made infrastructure within a 31-hectare section of the campus in 2016 and 2022. We investigated species-

specific nest heights and the host plant species chosen for nesting.

Results: A maximum of 86 active nests were identified, consisting of five species of five genera (Lestrimellita,

Partamona, Scaptotrigona, Tetragonisca, and Trigona). From 2016 to 2022, the stingless bee abundance increased

by 26.5 %, but the species composition remained the same. Tree cavities were the most attractive nesting loca-

tions, and their use increased within the sampling period. Overall nest survival was >64.3 % for the study period.

Conclusion: All bee species utilized a variety of tree species, but strangler figs (Ficus spp., Moraceae) were the

most important for nest construction. Nest height depended on the species and architecture. This work provides

a framework for future studies on tropical social bee communities in urban areas and offers valuable information

on their nesting biology in this habitat.

Keywords: Meliponini, Costa Rica, stingless bees, bee colony, trees, urbanization, Neotropics.

RESUMEN

Cambios temporales en la diversidad y abundancia de nidos de abejas

sin aguijón en un ambiente urbano

Introducción: La creciente urbanización ha puesto en peligro de extinción a muchas especies en una medida

desconocida. Las abejas sin aguijón (Apidae: Meliponini) son polinizadores muy importantes de plantas tropica-

les. Algunas especies están bien adaptadas a las áreas urbanas y utilizan estructuras artificiales para construir sus

nidos. En Costa Rica, hay 59 especies de abejas sin aguijón, pero nunca se ha realizado un recuento de su riqueza

y abundancia en áreas urbanas.

https://doi.org/10.15517/rev.biol.trop..v73iS2.64528

SUPPLEMENT

SECTION: ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64528, mayo 2025 (Publicado May. 15, 2025)

INTRODUCTION

Urban areas worldwide are expanding rap-

idly due to human population growth. Con-

tinuous urbanization transforms natural and

rural environments, leading to the formation

of urban centers (Aronson et al., 2015; Mont-

gomery, 2008). Species abundance and diversity

depend not only on the level of urbanization

but also on the taxonomic group (Blair, 1996;

Lewis et al., 2015; Lindenmayer et al., 2002).

Urban centers often exhibit low species diver-

sity due to altered biological composition and,

consequently, altered ecological relationships

between these species (Marzluff, 2001; Mar-

zluff, 2017).

In tropical regions, the largest group of

social bees are stingless bees (Apidae: Meli-

ponini), with approximately 550 described spe-

cies; they exhibit an eusocial lifestyle with

perennial colonies (Grüter, 2020). Monitoring

bee colonies facilitates long-term demographic

studies, and the results of these studies can

be applied in stingless beekeeping, known as

“meliponiculture” (Grüter, 2020). Stingless bees

are important pollinators and play crucial roles

in ecosystem maintenance and food availability

(Heard, 1999; Klein et al., 2018; Roubik, 2023;

Slaa et al., 2006). Several stingless bee species

have been shown to adapt very well to urban

conditions (Roubik, 2023; Velez-Ruiz et al.,

2013; Vieira et al., 2016). However, studies on

the impact of urbanization on bee diversity and

abundance are relatively rare but more impor-

tant than ever due to rapidly increasing urban-

ization (Solano-Gutiérrez & Otárola, 2025). To

maintain bee diversity and thus ecosystem ser-

vices, including pollination, bee conservation is

essential and can only be achieved by develop-

ing protection measures based on bee ecology

and knowledge of bee distribution.

In Costa Rica, the Central Valley repre-

sents the largest urban settlement, including

the capital city San José and other major cities,

forming a metropolitan area inhabited by 60%

of the country’s population (Madrigal-Solís et

al., 2019; Muñoz et al., 2021). Costa Rica has 59

species of stingless bees (Moure et al., 2007), of

which several are abundant in urbanized envi-

ronments, especially when trees are abundant,

but the species composition and dynamics have

never been analyzed in an urban ecosystem in

Central America.

In this study, we quantified the frequency

and species richness of stingless bee nests in

2016 and 2022 on a university campus within

an urbanized area. Additionally, bee host plants

and nesting sites were documented provid-

ing information on strategies for survival in

urban environments. We assessed whether bee

diversity and abundance changed between 2016

and 2022; we calculated a bee nest survival

Objetivo: Describir la composición y dinámica de la comunidad de abejas sociales en el campus de la Universidad

de Costa Rica en San José, durante un período de seis años.

Métodos: Buscamos sistemáticamente nidos (colonias activas) de abejas sin aguijón en árboles, edificios, paredes

y otra infraestructura artificial dentro de una sección de 31 hectáreas del campus en 2016 y 2022. Investigamos

las alturas de los nidos de cada especie y las especies de plantas elegidas para anidar.

Resultados: Se identificó un máximo de 86 nidos activos que pertenecían a cinco especies de cinco géneros

(Lestrimellita, Partamona, Scaptotrigona, Tetragonisca y Trigona). De 2016 a 2022, la abundancia de abejas sin

aguijón aumentó un 26.5 %, pero la composición de especies se mantuvo igual. Las cavidades de los árboles fueron

los lugares de anidación más atractivos y su uso aumentó durante el período de muestreo. La supervivencia de los

nidos fue >64.3 % durante el período de estudio.

Conclusión: Todas las especies de abejas utilizaron una variedad de especies de árboles, pero los higuerones

estranguladores (Ficus spp., Moraceae) fueron los más importantes para la construcción de nidos. La altura del

nido dependió de la especie y su arquitectura. Este trabajo proporciona un marco para futuros estudios sobre

comunidades sociales de abejas tropicales en áreas urbanas y ofrece información valiosa sobre su biología de

anidación en este hábitat.

Palabras clave: Meliponini, Costa Rica, abejas sin agujón, colonia de abejas, árboles, urbanización, Neotrópico.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64528, mayo 2025 (Publicado May. 15, 2025)

rate and determined whether there was a turn-

over in species using a specific nest location.

Thus, we identify host plant species and evalu-

ate nesting location preferences: substrate and

nesting height.

MATERIALS AND METHODS

Location: The study was conducted at

the Rodrigo Facio Campus of the University

of Costa Rica (UCR), which occupies approxi-

mately 97 hectares across three contiguous

sites. The Campus is located in Montes de Oca,

San José, Costa Rica (9°56’15”N 84°03’01”W;

1200 m a.s.l.). The climate is classified as tropi-

cal, with an annual mean precipitation of 1 868

mm (±358 SD) and an annual mean tempera-

ture of 20 °C (CIGEFI, 2025). The region’s dry

season is from December to April, and the

rainy season is between May and November

(Sáenz et al., 2007).

The censuses considered only the site

called Finca 1 (31 ha), the first one established

in 1956, with the oldest buildings and trees. The

other two campus sections experienced large

disturbances (infrastructure development) at

the time of the study, and their sampling was

logistically unfeasible. The population of the

campus is approximately 40 000 people (UCR,

2024). The sampled area has a high density of

buildings and infrastructure (58 % impervi-

ous surfaces and 42 % natural, seminatural, or

green areas); the area surrounding the campus

is highly urbanized (Fig. 1).

The flora of the campus is diverse, with

a variety of native versus introduced and wild

versus planted trees, shrubs, and herbaceous

plants, which are intensively managed. This

flora provides a variety of nesting and foraging

Fig. 1. Campus of the University of Costa Rica (only Finca 1 is shown) and the surrounding urban area of Montes de Oca, San

José. The yellow line delineates the sampled area. Within this polygon, gray stripes represent roads, and white lines delineate

trails, sidewalks, and parking lots. Source: OEPI, UCR; Bing aerial 2024, Microsoft Corporation.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64528, mayo 2025 (Publicado May. 15, 2025)

resources for wild bees. The campus center

hosts two hectares of regenerated forest area.

Data acquisition and visualization: Data

were collected in 2016 (September–Decem-

ber) and 2022 (December–March). The area

was searched systematically for stingless bee

colonies (active nests) on trees and man-made

infrastructure. Every tree and every man-made

element were visually examined from all sides,

from the ground to the top. The campus area

was run through several times by foot for each

survey. In this context, “trees” include any sec-

tion (internal or external) of living or dead

trees, remnant tree trunks, palms, or bamboo.

“Infrastructure” refers to anything man-made,

such as walls, buildings or fences. Since no

ground-nesting species have been reported on

the campus in the past nine years of sampling,

ground nests were excluded from searching.

Each tree species hosting a nest was identified,

and each nest was marked with coordinates

using GPS (GPSmap® Garmin CSX60), with an

average error of less than 2 meters. We analyzed

tree species data only for 2022 because the trees

used by bees in 2016 were also used in 2022.

In 2022, we first searched all preexisting

nests based on the survey conducted in 2016,

as well as every new location. The same search

protocol was used for both samplings. The

use of tags was avoided to prevent attracting

the attention of users of the campus, which

could negatively affect the nests. Finally, each

colony was initially identified at the species

level based on the nest entrance architecture

and voucher specimen collected from each

nest. Representative specimens were deposited

at the Museum of Zoology (MZUCR) of the

University of Costa Rica.

Additionally, we measured nest height to

identify species-specific preferences in nest

construction using a distance meter (Leica

Disto™ D2). To identify species-specific differ-

ences in nest height, a Dunn (1964) Kruskal‒

Wallis test was performed. Lestrimelitta mourei

was excluded because of its low nest count

(n = 2). The test was performed in R (https://

www.R-project.org/; Version: 2022.02.0+443).

RESULTS

Nest abundance and species diversity:

In total, 68 stingless bee nests (2.13 nests/ha)

were found on the Campus of the University of

Costa Rica in 2016, and 86 (2.69 nests/ha) were

found in 2022, representing a 26.5% increase

in abundance. In both surveys, the same five

different stingless bee species were found:

Partamona orizabaensis (Strand, 1919), Tetrag-

onisca angustula (Latreille, 1811), Scaptotrigona

subobscuripennis (Schwarz, 1951), Trigona cor-

vina (Cockerell, 1913), and Lestrimelitta mourei

(Oliveira and Marchi, 2005). The abundance of

four species increased over the six-year period

(Table 1). Partamona orizabaensis was the most

abundant species, comprising nearly half the

nests found during both samplings, followed

by Te. angustula. The cleptoparasitic L. mourei

was the least abundant species, with only one

nest in each sampling (Table 1). Nest survival

was high, ≥ 56% for every species, except for L.

Table 1

Frequency and percentage of eusocial bee nests according to year on the Campus Rodrigo Facio of the University of Costa

Rica and changes between censuses.

Species 2016 2022 Increase in

abundance (%)

Survival

2016-2022 (%)

Count %Count %

Partamona orizabaensis 39 57.35 42 48.84 7.69 56.40

Tetragonisca angustula 16 23.53 21 24.42 31.25 68.75

Scaptotrigona subobscuripennis 6 8.82 12 13.95 100 83.30

Trigona corvina 6 8.82 10 11.63 66.67 100

Lestrimelitta mourei 1 1.47 1 1.16 0 0

Total 68 100 86 100 26.47 64.70

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64528, mayo 2025 (Publicado May. 15, 2025)

mourei. Additionally, only one of the original

nests was reoccupied by a different species; one

S. subobscuripennis nest was formerly occupied

by P. orizabaensis.

Nesting location preferences: With one

exception, T. cor vina, all species built nests

in preexisting cavities in trees or man-made

structures. In both surveys, most bee nests were

found on trees (Fig. 2). Tetragonisca angustula

and P. orizabaensis were the only stingless bees

found nesting in infrastructure. The number

of Te. angustula nests built on infrastructure

doubled between the censuses, but those of P.

orizabaensis decreased to one third.

In 2022, bees inhabited 26 different tree

species (Table 2). Among them, 32.4% of all

nests were found on two native species of fig;

Ficus jimenezii hosted the most nests, includ-

ing all the bee species found in this census

(Table 2). Furthermore, in Ficus costaricana,

three different bee species were found (S.sub-

obscuripennis, Te. angustula, and T. corvina).

Taken together, Cupressus lusitanica and

Spathodea campanulata, two introduced tree

species, hosted approximately 22% of the nests

(Table 2).

Partamona orizabaensis was found in 16

different tree species, of which most of the nests

were built in F. jimenezii, S. campanulata, and

C. lusitanica. This bee species also nests abun-

dantly in human infrastructure (Fig. 2). Scap-

totrigona subobscuripennis nests were evenly

distributed among eight different tree species.

The same distribution could be observed for

Te. angustula, which was found with one nest

each in seven different tree species, except in F.

jimenezii, which had five nests. Trigona corvina

nests were also distributed among five different

tree species.

Nesting height: The average nesting height

of all stingless bee nests found in this survey

on the campus was 4.07 m above ground level.

For pairwise comparisons, nests from L. mourei

were excluded since only two nests were found.

The height of the nests differed significantly

between species (Kruskal‒Wallis test, χ2= 27.91,

Fig. 2. Percentage of stingless bee nests per sampling year according to species and nesting substrate (infrastructure or tree)

on the Campus Rodrigo Facio of the University of Costa Rica.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64528, mayo 2025 (Publicado May. 15, 2025)

P < 0.001). The average nest height of T. corvina

was 12.99 m, which was significantly higher

than that of all the other species (Dunn test,

P < 0.001). Average nest heights from P. o r i z a -

baensis, S. subobscuripennis and Te. angustula

were 3.42 m, 2.55 m, and 2.10 m respectively,

and did not differ significantly from each other

(Dunn test P > 0.1; Fig. 3).

DISCUSSION

Nest abundance and species diversity:

The abundance and diversity of bee species

depend on various biotic and abiotic factors.

On the Rodrigo Facio campus of the University

of Costa Rica, only five species of stingless bees

were found. The key factors influencing nesting

success in stingless bees include food source

availability, nest site availability, and human

alterations in the environment (Eltz et al., 2002;

Grüter, 2020; Roubik, 2023). Urban areas typi-

cally lack sufficient food resources, which is a

primary factor that reduces bee diversity (Wil-

son & Jamieson, 2019). The city of San José,

where the study area is located, has reduced

tree coverage and limited green spaces. These

factors drastically reduce food availability and

nesting spaces available for many bee species,

Table 2

Tree species used by stingless bees for nest construction, categorized by their origin.

Family Species Status Nest count Percentage of nests

per tree species

Moraceae Ficus jimenezii Native 19 26.8

Cupressaceae Cupressus lusitanica Introduced 8 11.3

Bignoniaceae Spathodea campanulata Introduced 7 9.9

Moraceae Ficus costaricana Native 4 5.6

Boraginaceae Cordia eriostigma Native 3 4.2

Myrtaceae Melaleuca quinquenervia Introduced 3 4.2

Arecaceae Roystonea regia Introduced 2 2.8

Bignoniaceae Tabebuia rosea Native 2 2.8

Poaceae Bambusa sp. Introduced 2 2.8

Cupressaceae Chamaecyparis sp. Introduced 1 1.4

Anacardiaceae Mangifera indica Introduced 1 1.4

Anacardiaceae Tapirira mexicana Native 1 1.4

Arecaceae Elaeis guineensis Introduced 1 1.4

Bignoniaceae Jacaranda mimosifolia Introduced 1 1.4

Casuarinaceae Casuarina equisetifolia Introduced 1 1.4

Clusiaceae Garcinia mangostana Introduced 1 1.4

Fabaceae Cojoba arborea Native 1 1.4

Fabaceae Erythrina poeppigiana Introduced 1 1.4

Lythraceae Lagerstroemia speciosa Introduced 1 1.4

Malvaceae Ceiba pentandra Native 1 1.4

Meliaceae Cedrela odorata Native 1 1.4

Moraceae Ficus elastica Introduced 1 1.4

Moraceae Ficus sp. Introduced 1 1.4

Myrtaceae Eucalyptus sp. Introduced 1 1.4

Sapindaceae Cupania glabra Native 1 1.4

Verbenaceae Citharexylum donnell-smithii Native 1 1.4

Dead trees 4 5.6

Total   71 100

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64528, mayo 2025 (Publicado May. 15, 2025)

further limiting their abundance and diversity

(Solano-Gutiérrez & Otárola, 2025). In con-

trast, El Rodeo, a forest patch outside San José

(800-1 000 msnm, 24 km W from the sampled

area), which is surrounded by rural areas, hosts

13 genera of stingless bees, such as Cepha-

lotrigona, Geotrigona, Melipona, Nannotrigona,

Oxytrigona, Plebeia, Scaura, Tetragona, and Tri-

gonisca, which are not present in the study area

(M. F. Otárola, unpublished data).

Interestingly, the overall number of nests

on the campus increased by >26% within six

years. One of the most important factors for bee

colonialization is the availability of nesting sites

and food sources (Grüter, 2020). Here this cor-

responds to tree cavity availability and suitable

infrastructure, such as walls with cavities (Han-

son et al., 2021). Changes in vegetation and

infrastructure cover between the censuses were

minimal and should not have influenced the

differences found (e.g., no new or dead nests

were related to new buildings or cut trees).

A reasonable explanation for the increase

in nest abundance could be the decreased

presence of humans during the COVID-19

pandemic. The campus restricted student

access from March 2020 until March 2022. The

decrease in human-induced stress may have

facilitated the increase in the number of nests.

The effect of disturbance can be calculated

from the yearly mortality rate of stingless bees.

Stingless bee nests often live for many years,

while the longevity of workers is relatively

short, one to several months, queens usually

live between 1 and 3 years and sometimes even

longer (Grüter, 2020). The annual mortality

rates per species in the sampled area range from

0–7.9% (excluding L. mourei), and the highest

values are from the species that use man-made

cavities on infrastructure. The mortality rates

reported here are lower than those reported in

other studies. Eltz et al. (2002) monitored Tri-

gona collina nests in undisturbed and managed

forests in Borneo. They calculated a yearly nest

mortality rate of 13.5–15.0%. Velez-Ruiz et al.

(2013) estimated that the nest mortality rate of

Te. angustula was lower than 10%. Slaa (2006)

reported that colony survival depends on the

bee species and location. This study found

that the annual mortality rate of Te. angustula

significantly depended on the habitat structure;

colonies in deforested areas (not urbanized)

lived three times longer than those inhabit-

ing forests. Our sampling protocol would not

detect cases in which a colony died but the nest

was subsequently reoccupied by the same spe-

cies, and mortality could be underestimated.

Nest substrate preference: Urban areas are

structurally complex, offering many spaces for

cavity-nesting bee species. Our findings suggest

that trees are preferred for nest construction

over artificial structures, with only two species

using artificial structures for nesting (see Fig.

2). Partamona orizabaensis demonstrated the

most generalist and adaptable nesting behavior,

utilizing 16 tree species and various forms of

Fig. 3. Stingless bee nest heights. Boxplots include 50% of

the data points; the bar indicates the median. The outliers

are shown.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64528, mayo 2025 (Publicado May. 15, 2025)

infrastructure. This adaptability is related to its

nest structure, as the species can build in semi

open cavities and create exposed nests, allow-

ing the colonization of locations unsuitable

for other species. Tetragonisca angustula, the

smallest bee in the survey with a small colony

size, can use small-scale cavities in both trees

and infrastructure. Tetragonisca angustula is

particularly abundant in urbanized areas across

the Neotropics (Aidar et al. 2013; Fierro et al.

2012; Velez-Ruiz et al. 2013; Vieira et al. 2016),

with some studies indicating higher abundance

in urban areas than in undisturbed environ-

ments (Fierro et al. 2012).

The other three stingless bee species have

specific requirements that limit the avail-

ability of suitable nesting sites. Scaptotrigona

subobscuripennis exclusively builds nests on

living trees and requires large, well-formed

cavities, which are less abundant. Trigona

corvina constructs massive, exposed nests in

canopy branches and do not require cavi-

ties. Undisturbed canopies are abundant at

the campus site, suggesting that the number

of T. cor vina colonies will likely increase in

the future, continuing the observed trend of a

67% increase in abundance during the sample

period. Lestrimelitta mourei also requires well-

formed cavities for nesting, but its biology as

an obligatory cleptobiont limits its abundance,

as this species is likely to maintain only small

populations in this area.

Nesting height: There are no comparative

data on nest-building heights between urban

and undisturbed areas. The nests of T. corvina

were the highest and largest among all the spe-

cies observed. Trigona corvina nests are fully

exposed in tree canopies, whereas other spe-

cies typically expose only their nest entrance

(Grüter et al., 2016). These nests can weigh up

to 100 kg (Roubik & Moreno Patiño, 2009) and

are particularly vulnerable to predators and

disturbances, which may explain the highly

aggressive behavior of T. cor vina (Grüter, 2020)

and their exceptionally high nesting height. In

contrast, the tree cavities used by other spe-

cies are located primarily in the lower parts

of trees, leading to relatively lower nesting

heights, indicating that nest-building height is

influenced by nest architecture and the avail-

ability of suitable nesting sites in urbanized

areas (Grüter et al., 2016).

Host‒plant interactions and their impli-

cations for conservation: In the 2022 survey,

stingless bees used 23 different plant species

for nest construction. Strangler figs (Ficus spp.)

were the most important trees due to their

growth habit. These trees begin as epiphytes,

with their roots growing downward and encir-

cling the host tree, eventually killing it (Schütt

& Lang, 2004). The decomposing host tree

leaves an empty space within the fig tree stem,

creating ideal nesting sites for stingless bees

(Grüter, 2020; Hanson et al., 2021). The utili-

zation of figs in tropical urban areas could be

a crucial strategy for providing natural nest-

ing sites for eusocial bees. For instance, Ficus

jimenezii hosted all the stingless bee species

found on the campus. Ficus costaricana also

harbored three species, but this species reaches

smaller sizes than F. jimenezii, which is a prob-

able reason for the lower number of colonies

using this species. Introduced species, such as

Spathodea campanulata and Cupressus lusitan-

ica, harbored many nests due to their high

number of cavities. However, each bee species

utilizes a wide range of tree species, indicating

that the availability of suitable cavities, rather

than specific tree species, is critical for nesting.

The campus of the University of Costa Rica

in San José was created on old pastures and cof-

fee plantations and has been reforested since its

foundation, allowing large and diverse trees to

be present today, including large fig trees that

require several decades to kill the host tree and

eliminate its trunk. This landscape contrasts

with the surrounding city, where few trees are

present and most of them are small in size,

precluding the existence of large natural cavi-

ties. Our results demonstrate that large stingless

bee populations can develop in urban environ-

ments with complex tree coverage, especially if

large trees are present. This information should

guide management strategies that prioritize

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64528, mayo 2025 (Publicado May. 15, 2025)

reforestation and the maintenance of large

urban trees to support the establishment and

maintenance of stingless bee colonies in urban

environments. Combined with the ongoing

planting of native ornamental plants that offer

food resources, this can transform tropical cit-

ies into truly bee-friendly spaces.

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.

ACKNOWLEDGEMENT

We would like to thank Alexa Morales for

her help in creating the map of the campus,

Mario Blanco and Rafael Acuña for plant iden-

tifications and Paul Hanson for his support with

this project. The DAAD funded J.K.’s academic

internship at the University of Costa Rica.

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