Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

Richness and abundance of non-flying terrestrial mammals

in an urban environment in the Neotropic

Diego Salas-Solano1 *; https://orcid.org/0000-0002-5990-785X

Luis Sandoval2, 3; https://orcid.org/0000-0002-0793-6747

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

Bernal Rodríguez-Herrera2, 3; https://orcid.org/0000-0001-8168-2442

1. Veragua Foundation. 70102, Limón, Costa Rica; dsalas@veraguarainforest.com

2. Escuela de Biología, Universidad de Costa Rica, San José, 2060, Costa Rica; gilbert.barrantes@gmail.com

3. Centro de Investigación en Biodiversidad y Ecología Tropical, Universidad de Costa Rica; bernal.rodriguez@ucr.ac.cr;

biosandoval@gmail.com

Received 30-VIII-2024. Corrected 24-II-2025. Accepted 04-III-2025.

ABSTRACT

Introduction: Urban green spaces are becoming increasingly important refuges for native fauna. In Costa

Rica, most of the human population is concentrated in the Central Valley, producing drastic changes in natural

ecosystems as urbanization increases and consequently reducing the natural habitats of multiple species. Urban

development generally decreases the native diversity of mammals.

Objective: We described the richness and abundance of medium-sized terrestrial mammals in fragments of

second growth and secondary forest vegetation immersed in an urban matrix.

Methods: We conducted a landscape analysis to measure the green spaces and urban land cover, and surveyed

terrestrial mammals using night baiting traps, diurnal counts in transects, camera traps, and occasional reports.

Results: We found six native and three introduced species of terrestrial mammals, representing less than 50% of

the medium-sized mammal diversity reported for the Central Valley of Costa Rica 40 years ago. The common

raccoon (Procyon lotor) and the variegated squirrel (Sciurus variegatoides) were the most abundant species, both

capable of exploiting urban habitats. Introduced species, especially free-ranging and feral cats (Felis catus), were

the most common species in the study site. Free-ranging introduced species may increase predation and disease

transmission. Fragmentation, isolation, and lack of connectivity caused by urbanization are likely affect the popu-

lations of sloth species (Bradypus variegatus and Choloepus hoffmanni) in the study site.

Conclusion: Maintaining natural and seminatural spaces and native vegetation is essential to conserve urban

biodiversity.

Keywords: medium-sized mammals, urban green spaces, fragments vegetation, urbanization, mammal diversity,

Neotropic.

RESUMEN

Riqueza y abundancia de mamíferos terrestres no voladores en un ambiente urbano del Neotrópico

Introducción: Los espacios verdes urbanos se están convirtiendo en un refugio cada vez más importante para

la fauna nativa de Costa Rica. En Costa Rica, la mayor parte de la población humana se concentra en el Valle

Central, consecuentemente, la urbanización reduce constantemente la vegetación remanente natural. En mamí-

feros, el desarrollo urbano generalmente disminuye la diversidad de especies nativas.

https://doi.org/10.15517/rev.biol.trop..v73iS2.64529

SUPPLEMENT

SECTION: ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64529, mayo 2025 (Publicado May. 15, 2025)

INTRODUCTION

The urbanscape is rapidly increasing in

response to human population growth world-

wide, which as significant impact on biodiver-

sity, particularly in hotspot areas (Alberti, 2005;

Brooks et al., 2002; Magura et al., 2010; Myers

et al., 2000). These changes result in natural

habitat loss, species introductions, and the

development of artificial environments unsuit-

able for most native species in most Neotropical

countries (Biamonte et al. 2011; Grimm et al.,

2008; Joyce 2006; McDonald et al., 2008; McK-

inney, 2008; Seto et al., 2012).

Within most urbanscapes there are often

small fragments of vegetation that vary in

shape, area, vegetation structure, and isola-

tion (e.g., forest patches, urban and periph-

eral parks, corridors) that allow some species

to survive and reproduce (Aronson et al., 2017;

Biamonte et al. 2011; González-García 2009;

Melles et al. 2003), as well as facilitate the colo-

nization of new species (Mayer & Sunde, 2020).

Therefore, understanding the abundance, dis-

tribution, and behavior of urban species in

remaining natural and seminatural habitats is

essential to promote conservation efforts and

develop sustainable urban planning (Aronson

et al., 2014; Luck, 2007; Marselle, et al. 2021;

Schneider et al., 2010; Seto et al., 2012).

Non-flying terrestrial mammals are one

of the groups most affected by urban develop-

ment and the fragmentation of natural habitats

(Haight et al., 2023; Pacifici et al., 2020). Many

species, such as large carnivores, require exten-

sive natural habitats to establish territories and

maintain viable populations. These carnivores,

are rare or extinct in natural areas transformed

by urban expansion (Moll et al., 2018; Orde-

ñana et al., 2010; Presley et al., 2019; Smith

et al., 2017). Fossorial mammals are another

group whose populations have been drastically

reduced or disappeared in urban areas due to

soil compaction and loss of food sources (How

& Dell, 2000). Introduced non-flying terrestrial

mammals (e.g., mice, rats, cats, and dogs) that

prey upon native species and compete for the

same resources also pose a serious threat to

native mammals in urban areas (Van Helden

et al., 2020).

Non-flying terrestrial mammals are an

important group in controlling populations of

other species in natural and some human-mod-

ified ecosystems (Moll et al., 2020). However,

many species in this group are highly suscep-

tible to human intervention in their habitats,

leading to their extinction or their populations

being drastically reduced (Dowding & Mur-

phy, 2001). Our objective in this study is to

evaluate the richness and relative abundance

Objetivo: Describimos la riqueza y abundancia de los mamíferos medianos terrestres en fragmentos de vegeta-

ción en regeneración y bosques secundarios inmersos en una matriz urbana.

Métodos: Realizamos un análisis de paisaje utilizando sistemas de información geográfica para medir los espacios

verdes y la cobertura urbana. El estudio de mamíferos terrestres incluyó trampeo nocturno, conteos diurnos en

transeptos, cámaras trampa y registros ocasionales.

Resultados: Encontramos seis especies nativas y tres especies introducidas de mamíferos terrestres, lo que repre-

senta menos de un 50% de la diversidad de mamíferos terrestre medianos reportada para el Valle Central hace 40

años. El mapache (Procyon lotor) y la ardilla (Sciurus variegatoides) común fueron las especies más abundantes,

ambas con la capacidad de aprovechar hábitats alterados por el desarrollo urbano. Las especies introducidas,

especialmente el gato doméstico, representan un problema por ser eficientes depredadores de la fauna nativa y

por transmitir enfermedades. Las especies de perezosos (Bradypus variegatus y Choloepus hoffmanni) en el sitio

de estudio se ven afectadas por el aislamiento de los fragmentos de vegetación y la falta de conectividad generada

por la urbanización.

Conclusión: Mantener los fragmentos de vegetación natural en las ciudades es primordial para la conservación

de la biodiversidad.

Palabras clave: mamíferos medianos, espacios verdes urbanos, fragmentos de vegetación, urbanización, diversi-

dad de mamíferos, Neotrópico.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

of non-flying terrestrial mammals within natu-

ral habitats in urban areas that vary in urban

development and vegetation cover (e.g., natural

habitats or gardens). If the extension of natural

habitats, in relation to the area covered by the

urbanscape matrix predicts the richness and

abundance of non-flying terrestrial mammals,

we expect the richness and abundance of these

mammals to be higher in larger and more natu-

ral fragments.

MATERIALS AND METHOD

Study site: We conducted this study on the

campus of the Universidad de Costa Rica from

August 2014 to March 2017. This is an urban

area in the northeastern section of the Costa

Rican Central Valley, San José province, Costa

Rica (9°54’N 84°03’W; 1200 m.a.s.l.). Tracts

of secondary premontane forest covered part

of the study area. These patches are remnants

of a much larger, nearly continuous forest

that extended to the highlands of the Central

Mountain range approximately 60–50 years ago

(Di Stéfano et al., 1996; Joyce, 2006; Nishida et

al., 2009). Today, the original native vegetation

is scarce and restricted to small fragments of

remnant riparian vegetation; other vegetation

is composed of isolated trees and gardens (Fig.

1 A, B; Sandoval et al., 2019).

The campus includes three separated sec-

tors: (1) Central Campus, occupying a total

area of 35 ha, which includes buildings, park-

ing areas, open recreational areas, and two

fragments of secondary forest, the Reserva

Ecologica Leonelo Oviedo (RELO), bordered

by Los Negritos stream and the Jardín Botánico

Orozco (JBO); (2) Ciudad de la Investigación,

covering 24.9 ha, which includes buildings,

parking lots, lawns, riparian vegetation, and

Fig. 1. A) Different cover types in the surroundings of the study site. B) Different cover types in the campus of Universidad

de Costa Rica, San José, Costa Rica. C) Diurnal transects and capture night trap sites (Jardín Botánico Orozco, Reserva

Ecológica Leonelo Oviedo, and Riparian Vegetation from left to right) in the three major sectors of Universidad de Costa

Rica, San José, Costa Rica.

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pastures along the Los Negritos stream; and

(3) Instalaciones Deportivas, covering 35.2 ha,

which includes buildings, lawns used for sports

and recreation, riparian vegetation, and spaces

covered by trees, bushes, and pastures, with the

Torres River at the northern border of this area

(Fig. 1B).

Landscape Analysis: We measured the

percentage of vegetation and urban cover in

the three sectors using satellite photos (reso-

lution 1:5000) from the Information System

of the Forest Resources of Costa Rica (2012).

We used ArcGIS version 10.5.3 to classify the

landscape into three categories: (1) vegetation

cover, which includes forest fragments, riparian

vegetation, and other spaces covered by trees;

(2) open green areas, such as grass, gardens,

and pastures; and (3) urban cover, including

buildings, roads, sidewalks, and other areas

covered by pavement.

Terrestrial mammal survey: We sampled

terrestrial mammals from August 2014 to

March 2017 using three methods: (1) baited

night traps, (2) diurnal counts along transects,

and (3) camera traps. We also included occa-

sional reports of sightings of terrestrial mam-

mals collected on campus during the study

and information on specimens deposited in

the Zoology Museum of the Universidad de

Costa Rica.

We placed baiting traps from August 2014

to December 2015 in three forest fragments

during nighttime sampling: (1) the Reserva

Ecológica Leonelo Oviedo (RELO), (2) the

Jardín Botánico Orozco (JBO), and (3) riparian

vegetation (RV) along Los Negritos stream (Fig.

1C). We placed 20 box traps (model 108, 25 x

30 x 81 cm; Tomahawk Live Trap Co., Toma-

hawk, Wisconsin) per trapping night using

commercial cat food as bait (Purina® Felix®).

The traps were placed 20 m apart. Traps were

set from 1600 to 0700 h and checked the next

day. We conducted 27 trapping nights (15

RELO, 8 JBO, and 4 RV). The maximum num-

ber of days between trapping nights was 91 and

the minimum was four (average = 16 days, SD =

24). We placed traps two consecutive nights on

three occasions and three consecutive nights on

one occasion. We measured the sampling effort

as trap nights (number of traps multiplied by

the number of trapping nights).

We conducted diurnal counts using bin-

oculars by walking at a steadily pace along

four 1-km-long transects from 06:00 to 08:00

am every two weeks. We surveyed each tran-

sect 24 times from April 2016 to March 2017.

Transects were located in three areas within

a matrix composed of forest fragments, open

green areas, and buildings (Fig. 1C). We estab-

lished transects in the three university sectors

as follows: two transects at the Instalaciones

Deportivas (ID1 and ID2), one at the Ciudad

de la Investigación (CI), and one at the Central

Campus (CC). During each survey, we counted

all mammals observed per transect. The fre-

quency of observations per species was calcu-

lated as the number of individuals observed

per transect.

We used ten camera traps (Bushnell Tro-

phy Cam HD) in RELO from March to April

2015. The camera-trap method is based on

identifying animal species using photographs

and videos taken by automatic cameras. Cam-

eras were placed 50 cm above the ground

between 20 m and 40 m from each other. We

did not consider for the analyses pictures or

videos of small mammal species (less than

1kg), because they are difficult to have a cor-

rect identification. We repeated this protocol in

JBO from June to July 2015. Cameras remained

active for 35 consecutive days in RELO and 23

days in JBO. The sampling effort represents

the number of trap cameras multiplied by the

number of active days (camera days).

We took pictures of the free-ranging and

feral cats (Felis catus) during night baiting traps,

diurnal counts in transects, and occasional

observations. We also used the records from

camera trap videos to identify and estimate the

number of free-ranging and feral cats on cam-

pus during the study period.

Data analysis: The frequency of capture

was defined as the number of individual per

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

species by trapping night, diurnal count tran-

sects, and camera trapping day. We estimated

relative abundance using two methods: (1)

the mean of individuals captured per trapping

night and (2) the mean of individuals observed

per transect for different species. Based on the

mean value of relative abundance, occasional

reports, and captures in camera traps, we clas-

sified the species into three abundance cat-

egories: rare (< 1 individual/survey, occasional

report of species, or rarely captured in camera

traps), uncommon (1–3 individuals/survey, or

occasional capture in camera trap), and com-

mon (> 3 individuals/survey).

We used the Morisita similarity index to

compare the mean species richness and abun-

dance across the four transects. This index

ranges from 0 to 1, where values close to

0 indicate no similarity between a pair of

transects, and values close to 1 indicate a

high similarity between a pair of transects. We

built a cluster tree with a single linkage and

the Morisita similarity values to represent the

transect relationship according to the species

richness and abundance. We also conducted a

non-metric multidimensional scaling (NMDS)

with Bray-Curtis distance to compare the com-

position of non-flying mammals’ communities

between transects. Differences in composition

among transects were tested with a one-way

PERMANOVA with 9999 permutations. We

used PAST (version 4.11; Øyvind Hammer,

Natural History Museum, University of Oslo,

Norway) for Morisita, cluster, NMDS, and

PERMANOVA analyses. Values are reported as

means ± SD.

RESULTS

Land use: The three sectors varied in the

area covered by each land type. The Central

Campus and Ciudad de la Investigación had

more urban cover than Instalaciones Deporti-

vas, which had more vegetation and open green

areas than the other two sites (Table 1). The

vegetation covers at Central Campus and Ciu-

dad de la Investigación was restricted mainly

to the riparian vegetation along Los Negritos

stream (Fig. 1). The Central Campus also pre-

sented natural remnants of secondary forest in

two reserves RELO and JBO (Fig. 1).

Richness and relative abundances of ter-

restrial mammals: We registered nine species,

including six native species and three intro-

duced species, from nine families of non-flying

terrestrial mammals, using the three sampling

methods (Fig. 2, Table 2). We captured raccoon

(Procyon lotor), common opossum (Didelphis

marsupialis) and feral cats (Felis catus) in an

effort of 524 trap nights (281 RELO, 165 JBO,

and 78 RV). The mean catch per night trap was

5.2 (SD = 2.6). The common raccoon was the

species with the highest average catch per night

with 3.1 ± 2.7 (mean ± SD) individuals/night,

followed by common opossum 1.4 ± 1.5 indi-

viduals/night, and feral cats 0.6 ± 0.9 (Table 3).

We recorded six non-flying mammal spe-

cies during diurnal counts. The mean number

of individuals observed per transect per day

was 9.1 ± 4.7. Variegated squirrels (Sciurus

variegatodes) were the most common species in

the four transects. We observed the three-toed

Table 1

Size (ha) and percentage occupied by the different habitats in the three major sectors of the Universidad de Costa Rica, San

José, Costa Rica.

Land cover Campus Central Ciudad de la Investigación Instalaciones Deportivas Total

Urban 25.1 (71.5%) 18.6 (75%) 5.6 (16%) 49.3 (51.8%)

Vegetation 10 (28.5) 4.5 (18%) 17.7 (50%) 32.2 (33.8%)

Open green areas 01.7 (7%) 12 (34%) 13.7 (14.4%)

Total 35,1 24.8 35.3 95.2

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Fig. 2. Terrestrial mammals found in the campus of Universidad de Costa Rica, San José, Costa Rica, August 2014-March

2017. A) Common Opossum Didelphis marsupialis. B) Two-toed sloth Choloepus hoffmanni (Photograph by Raquel Bone

Guzmán). C) Three-toed sloth Bradypus variegatus (Photograph by José Gabriel Barquero Jackson). D) Central American

least shrew Cryptotis orophilus. E) Variegated squirrel Sciurus variegatoides. F) Domestic cat Felis catus G) Domestic dog

Canis familiaris. H) Common raccoon Procyon lotor.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

sloth (Bradypus variegatus) and two-toed sloth

(Choloepus hoffmanni) only in Central Cam-

pus. The maximum number of sloths observed

during a diurnal count suggests a minimum

of seven individuals of three-toed sloth and

three individuals of two-toed sloth on Central

Campus during the study period. Feral cats

were observed in all three sectors studied. The

northern raccoons and feral dogs (Canis famil-

iaris) were observed only in Central Campus

(Table 4). The two transects in Instalaciones

Deportivas showed 100% similarity in species

richness and abundance (Fig. 3). Transects in

Intalaciones Deportivas were 99% similar to

Ciudad de la Investigación (Fig. 3). Central

Campus had a 93 % of similarity with transects

of the other sites (Fig. 3). The non-flying mam-

mals’ community composition varied between

transects (PERMANOVA: F = 7.29, P < 0.001,

Fig. 4). Central Campus differed from the

Table 2

Terrestrial mammals found in the campus of Universidad de Costa Rica, San José, Costa Rica, August 2014-March 2017. Type

of record includes: night trap (NT), diurnal counts (DC), camera trap (CT) and occasional report (OR).

Taxa Common name Type of record Abundance category

ORDER DIDELPHIMORPHIA

Family Didelphidae

Didelphis marsupialis Linnaeus, 1758

Common Opossum NT, CT Uncommon

ORDER PILOSA

Family Megalonychidae

Choloepus hoffmanni Peters, 1858

Two-toed sloth DC Rare

Family Bradypodidae

Bradypus variegatus Schinz, 1825

Three-toed sloth DC Uncommon

ORDER EULIPOTYPHLA

Family Soricidae

Cryptotis orophilus Allen, 1895

Central American least shrew OR Rare

ORDER RODENTIA

Family Sciuridae

Sciurus variegatoides Ogilby, 1839

Variegated squirrel DC, CT Common

Family Muridae

Rattus norvegicus Berkenhout, 

Norway rat CT Rare

ORDER CARNIVORA

Family Felidae

Felis catus Schreber, 1775

Domestic cat NT, DC, CT, OR Uncommon

Family Canidae

Canis familiaris Linnaeus 1758

Domestic dog DC, CT, OR Rare

Family Procyonidae

Procyon lotor Linnaeus, 1758

Common raccoon NT, DC, CT, OR Common

Table 3

Terrestrial mammal captures (mean ± SD per capture night) during night trap sessions in the Reserva Ecológica Leonelo

Oviedo (RELO), Jardín Botánico Orozco (JBO), and the Riparian vegetation (RP) on the campus of Universidad de

Costa Rica.

RELO JBO RV Total

Traps nights 281 165 78 524

Nights of capture 15 8 4 27

Didelphis marsupialis 29 (1.9 ± 1.4) 0 11 (2.5 ± 1.3) 40 (1.4 ± 1.5)

Procyon lotor 54 (3.6 ± 2.8) 28 (3.5 ± 2.8) 2 (0.5 ± 0.6) 84 (3.1 ± 2.7)

Felis catus 3 (0.2 ± 0.4) 10 (1.3 ± 1.2) 4 (1 ± 0.8) 17 (0.6 ± 0.9)

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64529, mayo 2025 (Publicado May. 15, 2025)

other three sites (pairwise comparisons, P <

0.001). However, the two transects of instalcio-

nes deportivas where 100% similar (P = 0.31),

forming a separate group, while the Ciudad de

la Investigación was different to Instalaciones

Deportivas 2 (P = 0.003), but similar to Instala-

ciones Deportivas 1 (P = 0.14).

We obtained a total of 653 videos of mam-

mals in 580 camera days and identified a total

of six species. The most commonly recorded

species was the common raccoon, followed

by the common opossum (recorded only in

the RELO) and feral cats. Norway rats (Rattus

norvegicus) variegated squirrels, and dogs were

rarely recorded using this method (Table 5).

We identified 24 free-ranging domestic

cats on night baiting traps, diurnal counts in

transects, camera trap videos and occasional

observations. We reported one Central Ameri-

can least shrew Cryptotis orophilus captured on

April 7, 2016, by a domestic cat near the Los

Negritos stream. We deposited the specimen

in the Museum of Zoology at the University of

Costa Rica (Voucher UCR4694).

DISCUSSION

Wild mammal populations are subject to

extreme pressures from habitat reduction and

rapid urban expansion (He et al., 2014). The

accelerated expansion of the urbanscape in

Table 4

Terrestrial mammals observed per diurnal counts (mean ± SD) in four transects (CC: Central Campus, CI: Ciudad de la

Investigación, ID1 and ID2: Instalaciones Deportivas 1 and 2 respectively) in the campus of Universidad de Costa Rica, San

José, Costa Rica.

Specie CC CI ID1 ID2 Total

Sciurus variegatoides 6.83 ± 3.07 6.46 ± 3.15 8.46 ± 4.88 10.79 ± 5.69 8.14 ± 4.61

Bradypus variegatus 1.79 ± 1.28 0000.45 ± 1.00

Choloepus hoffmanni 0.79 ± 0.72 0000.20 ± 0.49

Felis catus 0.54 ± 0.59 0.21 ± 0.51 0.08 ± 0.28 00.21 ± 0.46

Procyon lotor 0.08 ± 0.41 0000.02 ± 0.20

Canis familiaris 00.33 ± 1.13 0 0 0.08 ± 0.57

Fig. 3. Similarity of species richness and abundance average

between the four studied transects based on the Morisita

similarity index and single linkage in the cluster building.

Table 5

Numbers of terrestrial mammals recorded using camera

traps in two forest fragments Reserva Ecológica Leonelo

Oviedo (RELO) and Jardín Botánico Orozco (JBO) on the

campus of Universidad de Costa Rica.

RELO JBO Total

Camera days 350 230 580

Active days 35 23 58

Mammals 391 262 653

Didelphis marsupialis 122 0 122

Procyon lotor 182 252 434

Felis catus 77 9 86

Rattus norvegicus 8 0 8

Sciurus variegatoides 1 1 2

Canis familiaris 1 0 1

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

Costa Rica has led to the loss of a significant

portion of natural and semi-natural habitats

(e.g., coffee plantations and second-growth

areas), increasing the isolation of native vegeta-

tion. The rapid loss of natural habitats due to

urban expansion severely impacts native bio-

diversity (Grimm et al., 2008; He et al., 2014;

McDonald et al., 2013; Sushinsky et al., 2013).

A significant percentage of the vegetation in the

study area is protected in small, isolated forest

fragments. However, over 50% of the area has

been occupied by buildings, pavement, and

other urban structures. Increasing urbanization

negatively impacts animal diversity, as the rich-

ness and abundance of most species correlate

positively with vegetative complexity and plant

species richness (Aronson et al., 2017; McKin-

ney, 2002; Savard et al., 2000). In Costa Rica’s

Central Valley, urbanization increased by 72%

from 1973 to 2006 (Biamonte et al., 2011). Spe-

cifically, at the Universidad de Costa Rica, the

accelerated construction of multiple buildings

and the little interest in preserving the natural

and seminatural areas have further threatened

urban biodiversity.

The loss of natural habitat is one of the

main causes of biodiversity decline (Aronson

et al., 2017; Brooks et al., 2002; Fahrig, 2003;

He et al., 2014; Pimm & Raven, 2000; Seto et al.

2012). For mammals, species richness decreases

significantly in areas with extreme urbaniza-

tion such as central urban cores (McKinney,

2006). Wilson (1983) reported over 25 species

of medium-sized terrestrial mammals in the

urban areas of the Central Valley in the 1980s.

We registered fewer than ten species. Various

medium-sized mammals, such as monkeys,

anteaters, rabbits, porcupines, agoutis, weasels,

and skunks, reported years ago, were not found

in this study. Thus, the isolated and deterio-

rated fragments sampled in this study support

only a small fraction of the original fauna

(Aronson et al., 2014; Biamonte et al., 2011).

Variegated squirrels were very common in

the three sectors of the study site. Species in the

genus Sciurus are well-known for their ability to

adapt to urban habitats worldwide (Bonnington

et al., 2014; Jokimäki et al., 2017; La Morgia

et al., 2017; McCleery et al., 2008; McCleery,

2009; Parker et al., 2014; Rézouki et al., 2014).

Fig. 4. Non-metric multidimensional scaling comparing the community composition of the four studied transect sampled

during diurnal counts using Bray-Curtis distance.

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64529, mayo 2025 (Publicado May. 15, 2025)

The squirrels use native and introduced trees

for roosting and feeding, and exploit human-

provided resources such as food, buildings,

poles, and electrical wiring, allowing them to

adapt well to urban areas. The high abundance

of variegated squirrels may be attributed to

the diverse native and nonnative plants in the

study site (Di Stefano et al., 1996), and squirrels’

capacity to exploit the resources these plants

provide. The species was observed feeding on

seeds, flowers, fruits, leaves, and branches of

native and non-native trees and vines.

Urbanization alters ecosystem processes

through human activities, which can degrade

habitats but also increase the availability of

other resources, such as garbage containers

and urban structures used as food sources and

den sites (Prange & Gerht, 2004). Species that

can efficiently exploit these new resources may

occur at higher densities in cities compared

to rural or natural areas (Fedriani et al., 2001;

McKinney, 2002; Prange et al., 2003; Prange &

Gerht, 2004). This can lead to the dominance of

a few or even a single species in urbanized areas

(Prange & Gerht, 2004). Generalist species, with

broad dietary and habitat requirements, often

respond positively to anthropogenic resources

(McKinney, 2002). This included common rac-

coons, common opossums, and even squirrels,

which in the study site used similar habitats,

foods, and roosting sites (Ladine, 1997; Kis-

sell & Kennedy, 1992; Prange & Gerht, 2004;

Shirer & Fitch, 1970). These species are also

tolerant of fragmentation and human presence,

which facilitates their survival in urban habitats

(Prange & Gerht, 2004).

Similar to other studies, raccoons at the

study site appear highly efficient at exploit-

ing anthropogenic resources, leading to their

high abundance (Prange et al., 2003; Prange &

Gehrt, 2004). The common opossum is wide-

spread in the Neotropics and adapts well to

different conditions (Prange & Gehrt, 2004).

It thrives in forests and urban environments,

with a generalist diet that includes fruits, small

animals, and human food residuals (Barros

& de Aguiar Azevedo, 2014). However, com-

pared to opossums, raccoons’ larger body

size and greater dexterity may allow better

access to trash cans and dumpsters (Prange &

Gehrt, 2004). Common opossums were rarely

observed utilizing these resources in the study

site. Additionally, differential human responses

to specific mammal species (i.e., charismatic

species vs repulsive ones) can affect mammal

community structure.

Three of the nine species found at the study

site were introduced mammals, consistent with

patterns observed in other taxonomic groups

such as reptiles, amphibians, invertebrates and

plants (Duguay et al., 2007; Gibbon et al.,

2000; McKinney, 2008; Tait et al., 2005), where

the proportion of nonnative species increases

toward the urban core. Nonnative species may

have detrimental impacts on the native biota

through competition, predation, herbivory,

habitat alteration, and disease spread (Dowding

& Murphy 2001; Manchester & Bullock, 2000;

McKinney, 2008). Rats and free-ranging or

feral cats are particularly harmful (Longcore et

al., 2009; van Heezik et al., 2010). For example,

free-ranging and feral cats are major predators

of birds, small mammals, reptiles, amphibians,

and fish in cities (Jessup, 2004; Loss et al., 2013).

Cat abundance is negatively correlated with the

densities of some small mammals (Baker et al.,

2003; McCleery, 2010). In one study, 69% of

the prey items brought home by domestic cats

were mammals (Woods et al., 2003). Addition-

ally, rats and feral cats spread diseases to other

species and pose published health concerns

(Costa et al., 2015; Easterbrook et al., 2007;

Robertson, 2008).

Differences in non-flying terrestrial mam-

mal community composition among sectors

on the university campus may be a result

of the natural habitat patch size and vegeta-

tion composition (Markovchick-Nicholls et al.,

2008). Contrary to expectations that small and

distant patches support fewer species due to

limited species movement and fewer resources

(Brooks et al., 2002; Faeth et al., 2011; Marzluff,

2005; Watling & Donnelly, 2006), we recorded

the highest mammal richness on the Central

Campus site. The site has smaller vegetation

patches and is more isolated by urbanization.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

However, we observed the occurrence of two

sloth species, raccoons, and opossum. The lack

of connectivity between natural fragments and

the presence of large roads and buildings act as

physical barriers, isolating sloth populations on

the Central Campus and preventing them from

reaching other sites. Meanwhile, opossums and

raccoons benefit from older buildings, which

provide roosting sites such as ceilings and

holes. In contrast, the other two study sites

(Ciudad de la Investigación and Instalaciones

Deportivas) have newer buildings that do not

offer such roosting spaces.

Preserving patches of natural habitat with-

in urbanized landscapes is often advocated as

a method of conserving natural communities

(Aronson et al. 2017; Marzluff & Ewing, 2001;

Rivkin et al., 2019; Rudd et al., 2002; Watling

& Donnelly, 2006). Better integration of nature

in urban environments not only preserves bio-

diversity but also benefits human well-being.

Maintaining high local diversity in urban envi-

ronments increases contact with elements of

the natural world, contributing to the well-

being and quality of life because accelerates

recovery from stress, enhancing observational

skills, and reasoning abilities (Alvey, 2006;

Brown & Grant, 2005; Dearborn & Kark, 2010;

Horwitz et al., 2001; McPhearson et al., 2016;

Tzoulas et al., 2007). This study represents the

first description of medium-sized mammals in

the metropolitan area of Costa Rica. The find-

ings of this study emphasize the significance of

protecting and promoting the establishment of

biological corridors and forest patches in urban

areas for the conservation of native mammals.

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 Victor Madrigal and the Red

de Áreas Protegidas (RAP) of the Universidad

de Costa Rica for generating the maps. The

Universidad de Costa Rica for proving materi-

als and equipment for this research. We thank

Raquel Bone Guzmán and José Gabriel Barqu-

ero Jackson for providing photographs. LS was

supported by Vicerrectoría de Investigación

under project B6773.

REFERENCES

Alberti, M. (2005). The effects of urban patterns on

ecosystem function. International Regional

Science Review, 28(2), 168–192. https://doi.

org/10.1177/0160017605275160

Alvey, A. A. (2006). Promoting and preserving biodiver-

sity in the urban forest. Urban Forestry & Urban

Greening, 5(4), 195–201. https://doi.org/10.1016/j.

ufug.2006.09.003

Aronson, M. F., La Sorte, F. A., Nilon, C. H., Katti, M.,

Goddard, M. A., Lepczyk, C. A., Warren, P. S.,

Williams, N. S. G., Cilliers, S., Clarkson, B., Dobbs,

C., Dolan, R., Hedblom, M., Klotz, S., Kooijmans, J.

L., Kühn, I., MacGregor-Fors, I., McDonell, M., Mört-

berg, U., ... & Winter, M. (2014). A global analysis of

the impacts of urbanization on bird and plant diver-

sity reveals key anthropogenic drivers. Proceedings of

the Royal Society B, 281(1780), 20133330. https://doi.

org/10.1098/rspb.2013.3330

Aronson, M. F., Lepczyk, C. A., Evans, K. L., Goddard, M.

A., Lerman, S. B., MacIvor, J. S., Nilon., S. C., & Vargo,

T. (2017). Biodiversity in the city: key challenges for

urban green space management. Frontiers in Ecology

and the Environment, 15(4), 189–196. https://doi.

org/10.1002/fee.1480

Baker, P. J., Ansell, R. J., Dodds, P. A., Webber, C. E.,

& Harris, S. (2003). Factors affecting the distri-

bution of small mammals in an urban area.

Mammal Review, 33(1), 95–100. https://doi.

org/10.1046/j.1365-2907.2003.00003.x

Barros, F. B., & de Aguiar Azevedo, P. (2014). Common

opossum (Didelphis marsupialis Linnaeus, 1758):

food and medicine for people in the Amazon. Journal

of Ethnobiology and Ethnomedicine, 10(1), 65.

Biamonte, E., Sandoval, L., Chacón, E., & Barrantes, G.

(2011). Effect of urbanization on the avifauna in a

tropical metropolitan area. Landscape Ecology, 26(2),

183–194.

12 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64529, mayo 2025 (Publicado May. 15, 2025)

Bonnington, C., Gaston, K. J., & Evans, K. L. (2014).

Squirrels in suburbia: influence of urbanisation on

the occurrence and distribution of a common exotic

mammal. Urban Ecosystems, 17(2), 533–546.

Brooks, T. M., Mittermeier, R. A., Mittermeier, C. G., Da

Fonseca, G. A., Rylands, A. B., Konstant, W. R., Flick,

P., Pilgrim, P., Oldfield, S., Magin, G., & Hilton-Taylor,

C. (2002). Habitat loss and extinction in the hotspots

of biodiversity. Conservation Biology, 16(4), 909–923.

Brown, C., & Grant, M. (2005). Biodiversity and human

health: What role for nature in healthy urban plan-

ning? Built Environment, 31(4), 326–338.

Costa, F., Wunder Jr, E. A., De Oliveira, D., Bisht, V., Rodri-

gues, G., Reis, M. G., Ko, A. I., Begon, M., & Childs, J.

E. (2015). Patterns in Leptospira shedding in Norway

rats (Rattus norvegicus) from Brazilian slum com-

munities at high risk of disease transmission. PLoS

Neglected Tropical Diseases, 9(6), e0003819.

Dearborn, D. C., & Kark, S. (2010). Motivations for conser-

ving urban biodiversity. Conservation Biology, 24(2),

432–440.

Di Stéfano, J. F., Nielsen, V., Hoomans, J., & Fournier, L. A.

(1996). Regeneración de la vegetación arbórea en una

pequeña reserva forestal urbana del nivel premontano

húmedo, Costa Rica. Revista de Biología Tropical,

44(2A), 575–580.

Dowding, J. E., & Murphy, E. C. (2001). The impact of

predation by introduced mammals on endemic sho-

rebirds in New Zealand: a conservation perspective.

Biological Conservation, 99(1), 47–64.

Duguay, S., Eigenbrod, F., & Fahrig, L. (2007). Effects

of surrounding urbanization on non-native flora

in small forest patches. Landscape Ecology, 22(4),

589–599.

Easterbrook, J. D., Kaplan, J. B., Vanasco, N. B., Reeves, W.

K., Purcell, R. H., Kosoy, M. Y., Glass, G.E., Watson, J.,

& Klein, S. L. (2007). A survey of zoonotic pathogens

carried by Norway rats in Baltimore, Maryland, USA.

Epidemiology & Infection, 135(7), 1192–1199.

Faeth, S. H., Bang, C., & Saari, S. (2011). Urban biodiversi-

ty: patterns and mechanisms. Annals of the New York

Academy of Sciences, 1223(1), 69–81.

Fahrig, L. (2003). Effects of habitat fragmentation on bio-

diversity. Annual Review of Ecology, Evolution, and

Systematics, 34(1), 487–515.

Fedriani, J. M., Fuller, T. K., & Sauvajot, R. M. (2001). Does

availability of anthropogenic food enhance densities

of omnivorous mammals? An example with coyotes

in southern California. Ecography, 24(3), 325–331.

Gibbon, J. W., Scott, D. E., Ryan, T. J., Buhlmann, K. A.,

Tuberville, T. D., Metts, B. S., Greene, J. L., Mills,

T., Leiden, Y., Poppy, S., & Winne, C. T. (2000). The

Global Decline of Reptiles, Déjà Vu Amphibians:

Reptile species are declining on a global scale. Six

significant threats to reptile populations are habitat

loss and degradation, introduced invasive species,

environmental pollution, disease, unsustainable use,

and global climate change. BioScience, 50(8), 653–666.

González-García, A., Belliure, J., Gómez-Sal, A., & Dávila,

P. (2009). The role of urban greenspaces in fauna con-

servation: the case of the iguana Ctenosaura similis in

the ‘patios’ of León city, Nicaragua. Biodiversity and

Conservation, 18(7), 1909.

Grimm, N. B., Faeth, S. H., Golubiewski, N. E., Redman,

C. L., Wu, J., Bai, X., & Briggs, J. M. (2008). Global

change and the ecology of cities. Science, 319(5864),

756–760.

Haight, J. D. Hall, S. J. Fidino, M. Adalsteinsson, S. A.

Ahlers, A. A. Angstmann, J. Anthonysamy, W. J. B.

Biro, E. Collins, M. K. Dugelby, B. Gallo, T. Green, A.

M. Hartley, L. Jordan, M. J. Kay, C. A. M. Lehrer, E.

W. Long, R. A. MacDougall, B., Magle, S. B., … Lewis,

J. S. (2023). Urbanization, climate and species traits

shape mammal communities from local to continen-

tal scales. Nature Ecology & Evolution, 1–13.

He, C., Liu, Z., Tian, J., & Ma, Q. (2014). Urban expansion

dynamics and natural habitat loss in China: a mul-

tiscale landscape perspective. Global Change Biology,

20(9), 2886–2902.

Horwitz, P., Lindsay, M., & O’Connor, M. (2001). Biodiver-

sity, Endemism, Sense of Place, and Public Health:

Inter-relationships for Australian Inland Aquatic Sys-

tems. Ecosystem Health, 7(4), 253–265.

How, R. A., & Dell, J. (2000). Ground vertebrate fauna

of Perth’s vegetation remnants: impact of 170 years

of urbanization. Pacific Conservation Biology, 6(3),

198–217.

Jessup, D. A. (2004). The welfare of feral cats and wildlife.

Journal of the American Veterinary Medical Associa-

tion, 225(9), 1377–1383.

Jokimäki, J., Selonen, V., Lehikoinen, A., & Kaisanlahti-

Jokimäki, M. L. (2017). The role of urban habitats

in the abundance of red squirrels (Sciurus vulgaris,

L.) in Finland. Urban Forestry & Urban Greening, 27,

100–108.

Joyce, A. T. (2006). Land use change in Costa Rica: 1996–

2006, as influenced by social, economic, political, and

environmental factors. Litografía e imprenta LIL, S.A.,

San José, Costa Rica.

Kissell, R. E., & Kennedy, M. L. (1992). Ecologic rela-

tionships of co-occurring populations of opossums

(Didelphis virginiana) and raccoons (Procyon lotor)

in Tennessee. Journal of Mammalogy, 73(4), 808–813.

La Morgia, V., Paoloni, D., & Genovesi, P. (2017). Eradica-

ting the grey squirrel Sciurus carolinensis from urban

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73 (S2): e64529, mayo 2025 (Publicado May. 15, 2025)

areas: an innovative decision-making approach based

on lessons learnt in Italy. Pest Management Science,

73(2), 354–363.

Ladine, T. A. (1997). Activity patterns of co-occurring

populations of Virginia opossums (Didelphis vir-

giniana) and raccoons (Procyon lotor). Mammalia,

61(3), 345–354.

Longcore, T., Rich, C., & Sullivan, L. M. (2009). Critical

assessment of claims regarding management of feral

cats by trap–neuter–return. Conservation Biology,

23(4), 887–894.

Loss, S. R., Will, T., & Marra, P. P. (2013). The impact of

free-ranging domestic cats on wildlife of the United

States. Nature Communications, 4, 1396.

Luck, G. W. (2007). A review of the relationships between

human population density and biodiversity. Biological

Reviews, 82(4), 607–645.

Magura, T., Lövei, G. L., & Tóthmérész, B. (2010). Does

urbanization decrease diversity in ground beetle

(Carabidae) assemblages? Global Ecology and Biogeo-

graphy, 19(1), 16–26.

Manchester, S. J., & Bullock, J. M. (2000). The impacts of

non-native species on UK biodiversity and the effec-

tiveness of control. Journal of Applied Ecology, 37(5),

845–864.

Markovchick-Nicholls, L., Regan, H. M., Deutschman, D.

H., Widyanata, A., Martin, B., Noreke, L., & Ann

Hunt, T. (2008). Relationships between human dis-

turbance and wildlife land use in urban habitat frag-

ments. Conservation Biology, 22(1), 99–109.

Marselle, M. R., Lindley, S. J., Cook, P. A., & Bonn, A.

(2021). Biodiversity and health in the urban environ-

ment. Current Environmental Health Reports, 8(2),

146–156.

Marzluff, J. M. (2005). Island biogeography for an urba-

nizing world: how extinction and colonization may

determine biological diversity in human-dominated

landscapes. Urban Ecosystems, 8(2), 157–177.

Marzluff, J. M., & Ewing, K. (2001). Restoration of fragmen-

ted landscapes for the conservation of birds: a general

framework and specific recommendations for urba-

nizing landscapes. Restoration Ecology, 9(3), 280–292.

Mayer, M., & Sunde, P. (2020). Colonization and habitat

selection of a declining farmland species in urban

areas. Urban Ecosystems, 23, 543–554.

McCleery, R. A. (2009). Changes in fox squirrel anti-

predator behaviors across the urban–rural gradient.

Landscape Ecology, 24(4), 483.

McCleery, R. A. (2010). Urban mammals. Urban Ecosystem

Ecology, 87–102.

McCleery, R. A., Lopez, R. R., Silvy, N. J., & Gallant, D.

L. (2008). Fox squirrel survival in urban and rural

environments. The Journal of Wildlife Management,

72(1), 133–137.

McDonald, R. I., Kareiva, P., & Forman, R. T. (2008). The

implications of current and future urbanization for

global protected areas and biodiversity conservation.

Biological Conservation, 141(6), 1695–1703.

McDonald, B. C., Gentner, D. R., Goldstein, A. H., & Har-

ley, R. A. (2013). Long-term trends in motor vehicle

emissions in US urban areas. Environmental Science &

Technology , 47(17), 10022–10031.

McKinney, M. L. (2002). Urbanization, biodiversity, and

conservation: the impacts of urbanization on native

species are poorly studied, but educating a highly

urbanized human population about these impacts can

greatly improve species conservation in all ecosys-

tems. BioScience, 52(10), 883–890.

McKinney, M. L. (2006). Urbanization as a major cause

of biotic homogenization. Biological Conservation,

127(3), 247–260.

McKinney, M. L. (2008). Effects of urbanization on spe-

cies richness: a review of plants and animals. Urban

Ecosystems, 11(2), 161–176.

McPhearson, T., Pickett, S. T., Grimm, N. B., Niemelä, J.,

Alberti, M., Elmqvist, T., Weber, C., Haase, D., Breuse,

J., & Qureshi, S. (2016). Advancing urban ecology

toward a science of cities. BioScience, 66(3), 198–212.

doi.org/10.1093/biosci/biw002

Melles, S., Glenn, S., & Martin, K. (2003). Urban bird diver-

sity and landscape complexity: species–environment

associations along a multiscale habitat gradient. Con-

servation Ecology, 7(1).

Moll, R. J., Cepek, J. D., Lorch, P. D., Dennis, P. M., Robison,

T., Millspaugh, J. J., & Montgomery, R. A. (2018).

Humans and urban development mediate the sym-

patry of competing carnivores. Urban Ecosystems, 21,

765–778.

Moll, R. J., Cepek, J. D., Lorch, P. D., Dennis, P. M., Robison,

T., & Montgomery, R. A. (2020). At what spatial scale

(s) do mammals respond to urbanization?. Ecography,

43(2), 171–183.

Myers, N., Mittermeier, R. A., Mittermeier, C. G., Da Fon-

seca, G. A., & Kent, J. (2000). Biodiversity hotspots for

conservation priorities. Nature, 403(6772), 853.

Nishida, K., Nakamura, I., & Morales, C. O. (2009). Plants

and butterflies of a small urban preserve in the Cen-

tral Valley of Costa Rica. Revista de Biología Tropical,

57, 31–67.

Ordeñana, M. A., Crooks, K. R., Boydston, E. E., Fisher,

R. N., Lyren, L. M., Siudyla, S., Haas, C. D., Harris,

S., Hathaway, S. A., Turschak, G. M., Miles, A. K., &

14 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73 (S1): e64529, mayo 2025 (Publicado May. 15, 2025)

Van Vuren, D. H. (2010). Effects of urbanization on

carnivore species distribution and richness. Journal of

Mammalogy, 91(6), 1322–1331.

Pacifici, M., Rondinini, C., Rhodes, J. R., Burbidge, A. A.,

Cristiano, A., Watson, J. E., Woinarski, J. C. Z., & Di

Marco, M. (2020). Global correlates of range contrac-

tions and expansions in terrestrial mammals. Nature

Communications, 11(1), 2840.

Parker, T. S., Gonzales, S. K., & Nilon, C. H. (2014). Sea-

sonal comparisons of daily activity budgets of gray

squirrels (Sciurus carolinensis) in urban areas. Urban

Ecosystems, 17(4), 969–978.

Pimm, S. L., & Raven, P. (2000). Biodiversity: extinction by

numbers. Nature, 403(6772), 843–845.

Prange, S., Gehrt, S. D., & Wiggers, E. P. (2003). Demogra-

phic factors contributing to high raccoon densities

in urban landscapes. The Journal of Wildlife Manage-

ment, 324–333.

Prange, S., & Gehrt, S. D. (2004). Changes in mesopredator-

community structure in response to urbanization.

Canadian Journal of Zoology, 82(11), 1804–1817.

Presley, S. J., Cisneros, L. M., Klingbeil, B. T., & Willig, M.

R. (2019). Landscape ecology of mammals. Journal of

Mammalogy, 100(3), 1044–1068.

Rézouki, C., Dozières, A., Le Cœur, C., Thibault, S., Pisanu,

B., Chapuis, J. L., & Baudry, E. (2014). A viable popu-

lation of the European red squirrel in an urban park.

PloS one, 9(8), e105111.

Rivkin, L. R., Santangelo, J. S., Alberti, M., Aronson, M. F.,

de Keyzer, C. W., Diamond, S. E., Fortin, M. J., Fraxee,

L. J., Gorton, A. J., Hendry, A. P., Yang, L., Losos, J. B.,

Maclvor, S., Martin, R. A., McDonnell, M. J., Miles, L.

S., Munshi-Sur, J., Ness, R. W., Newman, A. E. M., ...

& Liu, Y. (2019). A roadmap for urban evolutionary

ecology. Evolutionary Applications, 12(3), 384–398.

DOI:10.1111/eva.12734

Robertson, S. A. (2008). A review of feral cat control. Jour-

nal of Feline Medicine and Surgery, 10(4), 366–375.

Rudd, H., Vala, J., & Schaefer, V. (2002). Importance of bac-

kyard habitat in a comprehensive biodiversity conser-

vation strategy: a connectivity analysis of urban green

spaces. Restoration Ecology, 10(2), 368–375.

Sandoval, L., Morales, C. O., Ramírez-Fernández, J. D.,

Hanson, P., Murillo-Hiller, L. R., & Barrantes, G.

(2019). The forgotten habitats in conservation: early

successional vegetation. Revista de Biología Tropical,

67(2), 36–52.

Savard, J. P. L., Clergeau, P., & Mennechez, G. (2000). Bio-

diversity concepts and urban ecosystems. Landscape

and Urban Planning, 48(3-4), 131–142.

Schneider, A., Friedl, M. A., & Potere, D. (2010). Mapping

global urban areas using MODIS 500-m data: New

methods and datasets based on ‘urban ecoregions’.

Remote Sensing of Environment, 114(8), 1733–1746.

Seto, K. C., Güneralp, B., & Hutyra, L. R. (2012). Global

forecasts of urban expansion to 2030 and direct

impacts on biodiversity and carbon pools. Procee-

dings of the National Academy of Sciences, 109(40),

16083–16088.

Shirer, H. W., & Fitch, H. S. (1970). Comparison from

radiotracking of movements and denning habits of

the raccoon, striped skunk, and opossum in northeas-

tern Kansas. Journal of Mammalogy, 51(3), 491–503.

Smith, J. A., Suraci, J. P., Clinchy, M., Crawford, A., Roberts,

D., Zanette, L. Y., & Wilmers, C. C. (2017). Fear of the

human ‘super predator’reduces feeding time in large

carnivores. Proceedings of the Royal Society B: Biologi-

cal Sciences, 284(1857), 20170433.

Sushinsky, J. R., Rhodes, J. R., Possingham, H. P., Gill, T. K.,

& Fuller, R. A. (2013). How should we grow cities to

minimize their biodiversity impacts?. Global Change

Biology, 19(2), 401–410.

Tait, C. J., Daniels, C. B., & Hill, R. S. (2005). Changes in

species assemblages within the Adelaide metropolitan

area, Australia, 1836–2002. Ecological Applications,

15(1), 346–359.

Tzoulas, K., Korpela, K., Venn, S., Yli-Pelkonen, V.,

Kaźmierczak, A., Niemela, J., & James, P. (2007).

Promoting ecosystem and human health in urban

areas using Green Infrastructure: A literature review.

Landscape and Urban Planning, 81(3), 167–178.

van Heezik, Y., Smyth, A., Adams, A., & Gordon, J. (2010).

Do domestic cats impose an unsustainable harvest

on urban bird populations?. Biological Conservation,

143(1), 121–130.

Van Helden, B. E., Close, P. G., Stewart, B. A., Speldewinde,

P. C., & Comer, S. J. (2020). An underrated habitat:

Residential gardens support similar mammal assem-

blages to urban remnant vegetation. Biological Con-

servation, 250, 108760.

Watling, J. I., & Donnelly, M. A. (2006). Fragments as

islands: a synthesis of faunal responses to habitat

patchiness. Conservation Biology, 20(4), 1016–1025.

Wilson, D. E. (1983). Checklist of Mammals of Costa Rica.

In D. H. Janzen. (Ed.). Costa Rican Natural History

(pp. 443–447). Univ. Chicago Press.

Woods, M., McDonald, R. A., & Harris, S. (2003). Preda-

tion of wildlife by domestic cats Felis catus in Great

Britain. Mammal Review, 33(2), 174–188.