Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e56527, enero-diciembre 2025 (Publicado Feb. 24, 2025)

Bird composition of a Neotropical city of Chiapas, México:

has a metacommunity structure?

Esteban Pineda-Diez de Bonilla1*; https://orcid.org/0000-0002-2643-1787

Karina A. Vázquez-Morales1; https://orcid.org/ 0009-0005-6199-6918

Ernesto Velázquez-Velázquez1; https://orcid.org/0000-0003-1884-0502

1. Museo de Zoología, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes de Chiapas, Ciudad Universitaria,

Libramiento Norte Poniente 1150, Colonia Lajas Maciel, Tuxtla Gutiérrez, Chiapas. CP 29039. México;

esteban.pineda@unicach.mx (*Correspondence), k.andrea.vazqm@gmail.com, ernesto.velazquez@unicach.mx

2. Laboratorio de Diversidad Acuática y Biogeografía, Instituto de Ciencias Biológicas, Universidad de Ciencias y Artes

de Chiapas (UNICACH), Ciudad Universitaria, Libramiento Norte Poniente 1150, Colonia Lajas Maciel, Tuxtla

Gutiérrez, Chiapas. CP 29039, México; wilfredo.matamoros@unicach.mx

Received 08-IX-2023. Corrected 15-X-2024. Accepted 06-II-2025.

ABSTRACT

Introduction: The birds in tropical cities are subject to changes in community composition and structure based

on landscape properties and habitat heterogeneity. Urban landscapes offer a set of fragmented habitats restricting

dispersion and promoting the metacommunity structure of birds.

Objective: To analyze the structure of an urban bird community with a meta-community perspective.

Methods: With a spatial array of 60 bird sampling points between May 2015 and February 2016, and estimations

of landscape metrics with a satellite image classification, we measured the coherence, turnover, and boundary

clumping of three subsets of birds to fit a metacommunity structure pattern and correlate with landscape proper-

ties of Tuxtla Gutierrez, a neotropical city with potential for bird conservation.

Results: The bird species composition comprises a dominant subset of exotic and synurbic species, another of

resident abundant species, and one of rare native species. The dominant species seem to be determined by a

Clementsian structure, associated with a bird community with a similar response to the environmental gradient.

Both dominant and rare species showed a nested structure associated with environmental filtering processes,

such as the amount of available habitat. The occurrence and distribution of abundant to rare species composition

were associated with the amount of natural vegetation cover, whereas the dominant species abundance with the

extension of urban coverage.

Conclusions: The bird species composition in urban landscapes of tropical dry forest regions can be structured

as a metacommunity in response to a gradient of vegetation fragmentation due to anthropogenic changes.

Synurbic and native species do not show a homogenization of diversity composition. To ensure the conditions

for bird diversity conservation in tropical urban landscapes, knowledge of the correlation of landscape elements

with community structure and habitat conditions in fragmented environments is needed.

Key words: bird diversity; Clementsian structure; heterogeneity; urban landscape; conservation.

https://doi.org/10.15517/rev.biol.trop..v73i1.56527

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73: e56527, enero-diciembre 2025 (Publicado Feb. 24, 2025)

INTRODUCTION

Cities represent highly modified environ-

ments where loss and fragmentation of animal

habitats lead to reduced species richness and

community homogenization at several scales

(Aronson et al., 2014; Clergeau et al., 2001;

Faeth et al., 2011). Furthermore, urban areas

undergo habitat transformation with negative

effects on community structure, especially for

birds, including species richness and changes

of species composition (Beninde et al., 2015),

at a landscape scale (Aronson et al., 2014; Bel-

locq et al., 2016), and this effect is exacerbated

in tropical regions (Leveau et al., 2017; Mac-

Gregor-Fors et al., 2021). Local communities

in urban areas have low alpha diversity because

native bird species have reduced occupancy or

abundance, and at a landscape scale, beta diver-

sity has high similarity of composition between

local communities, due to the dominance in

distribution and abundance of synurbic species

(Francis & Chadwick, 2011), whether they are

exotic or urban exploiters (MacGregor-Fors

et al., 2010; Martin-Etchegaray et al., 2018).

Conversely, tolerant native species occurring in

cities can increase the local community dissimi-

larity by restricted dispersion between remnant

fragments of habitat on urban landscapes thus

bird community structure and species compo-

sition change (Clergeau et al. 2001; Escobar-

Ibañez et al. 2020).

The community structure of species from

a metacommunity perspective can be described

based on abundance and distribution patterns

of the species, within dispersal restricted land-

scapes as the disturbed heterogeneous or urban

gradient landscapes (Alberti & Wang, 2022).

The distribution of three components, coher-

ence as the degree of spatial continuity of

species across a major gradient; turnover as

the species replacement from site to site and

RESUMEN

Composición de aves de una ciudad Neotropical de Chiapas, México:

¿Tiene una estructura metacomunitaria?

Introducción: Las comunidades de aves en las ciudades tropicales están sujetas a cambios en su composición

y estructura según las propiedades del paisaje y la heterogeneidad del hábitat. El paisaje urbano ofrece hábitats

fragmentados que restringen la dispersión y promueve la estructura metacomunitaria de las aves.

Objetivo: Analizar la estructura de la comunidad de aves urbanas con un enfoque de arreglos metacomunitarios.

Métodos: Con una matriz espacial de 60 puntos de muestreo de aves entre mayo de 2015 y febrero de 2016, y

estimaciones de métricas del paisaje con una clasificación de imágenes satelitales, medimos la coherencia, el

recambio y el agrupamiento de tres subconjuntos de aves para ajustarlos a un patrón de estructura metacomunita-

ria y correlacionarlas con las propiedades paisajísticas de Tuxtla Gutiérrez, una ciudad Neotropical, con potencial

para la conservación de las aves.

Resultados: La composición de aves es un subconjunto de especies dominantes exóticas y sinúrbicas, uno de

especies residentes abundantes y uno de especies raras nativas. Las especies dominantes están determinadas por

una estructura Clementsiana, asociada a una comunidad de aves con una respuesta similar al gradiente ambiental,

mientras que tanto especies dominantes como raras muestran una estructura anidada, asociado con procesos

de filtro ambiental como la cantidad de hábitat disponible. La ocurrencia y distribución de la composición de

especies abundantes y raras se asocia con la cantidad de cobertura vegetal natural, mientras que la abundancia de

especies dominantes con la extensión de la cobertura urbana.

Conclusiones: La composición de especies de paisajes urbanos en regiones tropicales secas puede tener una

estructura metacommunitaria en respuesta a un gradiente de fragmentación de la vegetación por cambios

antropogénicos. Las especies sinúrbicas y nativas no muestran una homogeneización de la composición de la

diversidad de aves. Para asegurar las condiciones de la conservación de aves en los paisajes tropicales urbanos,

es necesario conocer la estructura de la comunidad y los elementos paisajísticos que ofrecen las condiciones de

hábitat en ambientes fragmentados.

Palabras clave: diversidad de aves; estructura Clementsiana; heterogeneidad; paisaje urbano; conservación.

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boundary clumping as the extent to which

the limits of ranges of the different species are

clustered together produce at least, six pos-

sible idealized composition patterns (Leibold

& Mikelson, 2002; Presley et al., 2010). These

vary, from a random arrangement (Gleasonian

pattern) to an arrangement of sites grouped by

a composition of similar species (Clementsian

pattern, Presley et al., 2010). Based on this

spatial distribution arrangement of species of a

metacommunity in a landscape can be related to

an underlying emergent process, such as species

sorting or mass effect (Holyoak et al., 2005; Lei-

bold et al., 2004). Bird community composition

has been described in urban and non-urban

settings (Beninde et al., 2015; MacGregor-Fors

& Schondube, 2012). However, studies focusing

on bird metacommunity structure in an urban

landscape gradient has been very little explored

(Ferenc et al., 2014).

Habitat reduction has been identified as the

main driver of native bird richness reduction in

urban settings. Urbanization, typically reduces

the proportion of vegetation cover compared to

build area and consequently reduces the distri-

bution and abundance of most sensitive native

bird species (Clergeau et al., 2001; Escobar-

Ibañez et al., 2020). Heterogeneity of land cover

types and configuration inside urban landscape

can soften the filtering effect of urban cover

for these birds and may be a major determi-

nant for species distribution (Lepczyk et al.,

2017), and community structure (Beninde et

al., 2015; Silva et al., 2016). The dominance of

synurbic bird species seems to be regulated by

the extension of non-vegetative covers and the

heterogeneity of urban landscapes (Silva et al.,

2016). Those aspects of the urban landscape

have not been studied for bird communities in

the Neotropical region (Bellocq et al., 2016),

where there are many endemic birds (Gordon

& Ornelas, 2000).

Under the metacommunity perspective,

beta diversity properties like nestedness, turn-

over and similarity allow identification of the

metacommunity structure of species composi-

tion in heterogeneous landscapes or environ-

mental gradients. In particular urbanization

is expected to produce a random composition

of synurbic species with a low beta diversity

due to the effect of homogenization (Presley et

al., 2010; Wang et al., 2019), a nested structure

of native species between local communities

(Baselga, 2010), and a positive coherence and

low local (alfa) diversity (Presley et al., 2010;

Wang et al., 2019). These patterns are due to

abundance change of native species with syn-

urbic dominant species (Francis & Chadwick,

2011). Likewise, under urbanization diversity

partitioning patterns predict low beta diversity

and a small difference between range of alfa and

gamma diversity (Wang et al., 2019). The aim of

this study is to determine whether there was a

homogenized (Gleasonian pattern) or discrete

subset of species (Clementsian pattern) com-

position of land bird in the urban landscape on

the tropical city of Tuxtla Gutiérrez, Chiapas,

Mexico, and determine what land cover of

urban landscape are related to the occurrence

and abundance of dominant, abundant and rare

bird species.

MATERIALS AND METHODS

Study area: The study was carried out

in Tuxtla Gutiérrez city (TG), the largest city

and capital of the Southern state of Chiapas in

México. It covers around 83.5 km2 (Silva et al.,

2015), and is located at 600-750 m.a.s.l. Tuxtla

Gutiérrez is surrounded by hills with remnants

of preserved natural vegetation, some of these

fragments contain a surrounding corridor of

protected areas (Altamirano-González-Ortega

et al., 2007). The original vegetation that sur-

rounds the city is tropical dry forest, which is

distributed along the Central Valley region of

Chiapas. Small remnants of this vegetation are

scattered within the city with different succes-

sional phases along the banks of the Sabinal

River basin which crosses the city from West

to east, (Díaz-Pascacio et al., 2018). The urban

area is dominated by buildings, pavement and

dirt roads, the vegetation cover within the

city is currently reduced and fragmented to

only 10 % of the original surface (Silva et al.,

2015). The landscape is heterogeneous, with

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altered vegetation covers represented by urban

parks, gardens, wastelands and some remain-

ing agricultural plots. On the outskirts of the

city, there are tree vegetation fragments, such

as suburban, parks and reserves, as “Cañon

del Sumidero” National Park which represent a

regional reservoirs of bird diversity (Altamira-

no-González-Ortega et al., 2007).

Bird sampling design: Sixty bird point

count locations were established along the

heterogeneous urban landscape on TG, in

which the different types of land covers were

estimated (Fig. 1). There was a minimum dis-

tance of 600 m between each point to represent

the heterogeneity of the city’s landscape and

maintain the independence of the bird records,

and maximize sample size (Buckland, 2006).

The point counts had a fixed radius of 25 m

(Ralph et al., 1993), and all the land birds seen

or heard were recorded during a time of 10

min, early in the morning between 6:30 and

Fig. 1. A. Distribution of bird sampling points centered in (black circles) areas of estimated land cover types within Tuxtla

Gutiérrez urban landscape. Vegetation covers in green shades and urban covers in gray shades. B. location of Chiapas state

in Southern México. C. location of Tuxtla Gutiérrez urban landscape in the Chiapas state.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73: e56527, enero-diciembre 2025 (Publicado Feb. 24, 2025)

9:30 h. Each point was visited on four occa-

sions between May 2015 and February 2016.

With this sampling method, average of relative

abundance (ab) and species richness (r) was

estimated for all resident land birds. Migratory

birds were excluded from the analysis, as well

as birds that fly over the vegetation of land

city covers, such as swifts (Apodidae), swal-

lows (Hirundinidae) and vultures (Cathartidae)

because they do not use the urban landscape

(Pineda-Diez de Bonilla et al. 2012).

Urban landscape description: To describe

the landscape properties of TG, we estimated

the amount of seven land cover classes within a

circular area of 280 m radius (24.6 ha), associ-

ated with each point count. Seven class covers

were used in an unsupervised classification

with an ISODATA clustering of a Landsat 8

satellite image of 30 m resolution provided by

U.S. Geological Survey (2016) from November

2015 (Fig. 1). This classification defines limits

between different types of natural and artificial

land covers, based on the vegetation structure

and density, human bare ground covers were

defined, based on previous experience of sur-

veys, these covers can be extrapolated to the

entire urban area of the study landscape (Feeley

et al., 2007). The area of each cover class is

within 24.6 ha. circular area was estimated in

QGIS 2.8 (QGIS Development Team, 2015)

(Fig. 1). To describe the landscape properties

commonly related with specie richness and

diversity in disturbed landscapes (Beninde et

al., 2015; Rojas et al., 2017), nine landscape

indices were estimated from the area amount of

the land cover classes with the Fragstat program

(McGarigal & Marks, 1995), These were, the

number of patches (NP), class richness (PR),

the habitat heterogeneity index (SHDI), the

landscape fragmentation with the Intersper-

sion and juxtaposition index (IJI), the cover-

age of largest patch index (LPI), landscape

shape index (LSI), mean perimeter- area ratio

(PARA), fractal dimension irregularity index

of perimeter- area ratio (PAFRAC), and class

fragmentation index (MESH) was calculated.

Bird species subsets: To analyze separately

the effect of synurbic and native species on the

structure of the urban bird metacommunity,

we divided the total bird species registered into

dominant (D), abundant (A) and rare (R) spe-

cies, based on similarity-based entropy (Grab-

chak et al., 2017). Then, we used Hill numbers

(Chiu & Chao, 2014) to decompose the effec-

tive species richness into the three exponent

values of Q (0, 1, 2). These numbers allow us to

separate the number of dominant species (Q2),

abundant species (Q1-Q2) and rare species (Q0-

Q1) from total species (Q0) in the metacommu-

nity. Additionally, with this partitioning scheme

we estimate the relative contribution of relative

species abundance of local community assem-

blage (alpha diversity), into the metacommuni-

ty assemblage (gamma diversity), between local

assemblages (beta diversity) or effective num-

ber of local communities (Marcon & Hérault,

2015). For these purposes of decomposition of

diversity entropy, we used the DivPart function

of the entropart package (Marcon & Herault,

2015) in the R software (R Core Team, 2015).

Metacommunity structure: To determine

the bird metacommunity structure, and the

change in the idealized structure for the three

species subset (dominant, abundant, rare spe-

cies), we analyzed the coherence, turnover and

boundary clumping and their fit to a meta-

community structure (Dallas, 2014; Leibold &

Mikkelson, 2002; Presley et al., 2010). A species

by sites matrix was arranged for each spe-

cies subset and total species metacommunity.

Coherence was estimated testing the number

of embedded absences in the ordered matrix

of species by sampling locations, a coherence

analysis was used for each subset species group.

Turnover analysis was evaluated to compare

the number of observed replacements of paired

species along sites, the observed replacements

is compared against to a random null model

patterns, to pick out between a negative sig-

nificance (nestedness), positive significance

(clumped) or not significance (quasi-struc-

ture), subsequently boundary clumping analy-

sis was tested for a positive significance to

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Clementsian structure, negative significance to

Gleasonian structure and evenly spaced struc-

ture for non-significance arrange. Boundary

clumping was evaluated by a goodness of fit test

of the observed distribution with the expected

equiprobable distribution of range boundaries

of species with the Morisita’s index. All these

metacommunity measures were analyzed in R

(R Core Team, 2015) with the use of the meta-

com (Dallas, 2014) package.

Correlation of bird and urban landscape

variables: To determine the association of the

species of each group with cover types and

landscape indices variables, we performed

a redundancy analysis (RDA). This analysis

tested the association of area land cover and

landscape indexes to each group of land bird

species and determined the influence of urban

landscape variables on the presence and struc-

ture of bird species community (Legendre &

Anderson, 1999). We implemented the RDA

procedure in the R package vegan (Oksanen et

al., 2007) in R (R Core Team, 2015).

RESULTS

We recorded an overall of 4 006 individuals

and 63 bird species, along the 60-point counts.

We found four exotic species present in most

of the urbanized sites (Rock Dove Columba

livia, House Sparrow Passer domesticus, Eur-

asian Collared-dove Streptopelia decaocto and

Monk Parakeet Myiopsitta monachus), and four

native species considered synurbic to Mexico

(Common Ground-dove Columbina passeri-

na, Great-tailed Grackle Quiscalus mexicanus,

Clay-colored Thrush Turdus grayi and Tropi-

cal Kingbird Tyrannus melancholicus), most of

these species belonged to the subset of domi-

nant species. The remaining bird species were

native species present in the mosaic of tropical

dry forest, as the natural vegetation in the

region (Fig. 2).

According to expected species richness

a completeness of 89 % was of the maximum

number of expected species estimated with

the Clench accumulation model (Soberón &

Llorente, 1993). This result suggested that our

sampling effort represents the compositional

pattern of dominant, abundant and rare bird

species presence occurring in TG. Land cover

distribution in the urban landscape of TG

shows an urbanization gradient pattern (Fig. 3),

represented by a total average extent of 35 % of

urban cover (n = 56), 22 % for herbaceous (n

= 60) and 20 % for bare ground cover (n = 54)

respectively, and an average richness cover of

five cover types for all sampling points.

According to the diversity decomposition

based on entropy, bird metacommunity of TG

has an average of 10.25 effective richness value

to local communities as alpha diversity, and

6.14 effective local communities in the meta-

community as beta diversity with 63 species

richness to gamma diversity. This result indi-

cates that spatial scale used in our study to

describe the metacommunity structure of birds

in the urban landscape seems to be adequate,

according to the effective number of local com-

munities estimated with the decomposition of

diversity (Marcon & Hérault, 2015) and homo-

geneous distribution across the urban land-

scape of the commonly considered as urban

exploiters species (Table 1).

Table 1

Diversity partition in effective number of total species (Q0),

abundant species (Q1) and dominant species (Q2), for alpha,

beta and gamma components of the bird metacommunity

of the Tuxtla Gutiérrez city of Chiapas.

Diversity Q0Q1Q2

Alpha 10.25 7.34 5.61

Beta 6.14 3.05 2.55

Gamma 63 22.44 14.34

We use the results to the bird metacom-

munity structure analysis to fit one of the

metacommunity structure models to the birds

of TG. We found a variation on the urban land

bird metacommunity structure within each

species subset, the total bird metacommu-

nity and dominant subset has a Clementsian

structure, whereas abundant and rare species

subsets shows a nestedness structure (Table 2).

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Fig. 2. Rank abundance distribution of the 63 land bird species metacommunity recorded in the landscape of Tuxtla

Gutierrez city.

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Dominant species subset of bird was made of

native opportunistic species and exotic species,

and according to their spread distribution in

the point counts and two groups of birds were

formed, the rare species group composition

represents terrestrial native species present on

the TDF vegetation.

The association of each subset bird spe-

cies to urban landscape properties in the RDA

showed different relationships for dominant,

Table 2

Elements of a metacommunity structure pattern of land birds in the urban landscape of Tuxtla Gutiérrez city, Chiapas.

Elements E. Abs Total Dominants Abundant Rare

Coherence E. Abs 1.54E + 03 3.49E + 02 1.44E + 02 7.56E + 00

P1.40E - 09 3.57E - 17 5.99E - 07 1.30E - 01

Mean 2.41E + 03 4.24E + 02 2.12E + 02 8.26E + 02

SD 3.94E + 01 8.90E + 00 1.37E + 01 4.60E + 01

Turnover Rep 2.31E + 05 6.85E + 03 1.31E + 03 1.49E + 03

P6.75E - 01 1.59E - 03 1.62E - 06 5.06E - 02

Mean 2.43E + 05 4.37E + 03 3.34E + 03 1.62E + 04

SD 2.98E + 04 7.88E + 02 4.14E + 02 1.40E + 03

Boundary Clumping Morisita Index 5.23 1.79 1.86 1.87

P 0.000 5.00E-10 1.73E - 13 3.40E - 08

Df 60 12 6 38

Interpretation Clementsian Clementsian Nestedness Nestedness

E. Abs: Observed embedded absences; Rep: Number observed replacements; P: Probability test; Mean: Estimated random

values; SD: Standard deviation; Df: degree freedom.

Fig. 3. Distribution of % cover and heterogeneity index (SHDI) values of the seven cover types of 60 sampling sites inside the

urban landscape of Tuxtla Gutierrez city.

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abundant and rare species. Interspersion and

juxtaposition index (IJI), heterogeneity index

(SHDI) and mean perimeter area ratio (PARA)

were the most important landscape index, and

amount of cover area of trees, shrubs, herbs and

urban were the most important land cover for

dominant bird species (Fig. 4).

DISCUSSION

The structure of bird metacommunity and

spread distribution of native and synurbic bird

species in tropical urban landscapes has been

poorly described (Martin & Bonier, 2018). Bird

species composition of TG city resembling

a metacommunity like vertebrates in other

cities studied in Mexico (e.g., Charre et al.,

2013; Escobar-Ibañez et al., 2020). However, a

homogenization effect by the dominant spe-

cies and a lower diversity was found in sites

with predominant urban cover, it was similarly

to urban mosaic in many cities (Leveau et al.,

2017; Marzluff, 2016), where exotic and syn-

urbic species dominate the metacommunity

structure.

The results of decomposition of total bird

diversity according to the entropy analysis

between sampling sites (alpha diversity), sug-

gest that the metacommunity is composed of

6.14 effective local communities. A metacom-

munity structure is maintained even when we

discard the subset of rare species, reducing the

structure to three local communities. This is

due on the one hand, to the fact that rare spe-

cies with restricted distribution in the urban

landscape tend to increase dissimilarity and

gamma diversity within the city of TG. On the

other hand, the reduction in the number of

local communities is due to unlimited dispersal

capacity of the dominant species with a general-

ist habitat use, and present in different habitat

types, such as the ground pigeon Columbina

passerina, the large-tailed grackle Quiscalus

mexicanus or the wood thrush Turdus grayi,

which increase the similarity between local

communities (MacGregor-Fors et al., 2021).

This result of bird community composition in

the urban landscape of TG is consistent with

a not homogeneous effect of environmental

filtering in bird communities (Silva et al., 2016).

Two different metacommunity structures were

found with the analysis of coherence, turnover

and boundary clustering, between the subset

species groups and the association within the

environmental gradient. A Clementsian struc-

ture (Presley et al., 2012) for both the total num-

ber of birds and the subset of dominant species,

this structure is associated with local communi-

ties with a discrete and coherent replacement

of a species subset composition between local

communities or sites, in response to distribu-

tion processes of similar environmental factors

along the urban gradient (Leibold et al., 2004).

This metacommunity structure coincides with

the structure of other Neotropical vertebrates

(López-González et al., 2012; Ochoa-Ochoa &

Whittaker, 2014). A nested metacommunity

structure shown by both the abundant and rare

subset species, is associated to a loss of distribu-

tion of habitat specialized species within a nar-

row gradient of habitat availability due to the

size and extent of fragments of vegetation cover

within the urban landscape (Gómez-Moreno

et al., 2023; Henriques-Silva et al., 2013), all

abundant and rare species are native residents

of tropical dry forest surrounding of TG.

The affinity for a different type of struc-

ture is due to differences in the distribution of

species between sites, which show an impor-

tant gradient of the urban landscape, which

describes the increase in the extent of urban

coverage and the decrease in tree vegetation.

and by the gradient of coverage heterogeneity

in the landscape (Fig. 3). The positive affin-

ity of the dominant species subset with urban

cover and habitat heterogeneity is consistent

with the results of other studies that evaluate

the relationship of habitat quantity hypotheses

(Beninde et al., 2015) and habitat diversity

(Mateo-Tomás et al., 2019). This is due to the

widespread distribution pattern, the abundance

of these synurbic species (MacGregor-Fors et

al., 2010; Martin-Etchegaray et al., 2018) and

the fact that they may have behavioral adjust-

ment to the urban environment (MacGregor-

Fors et al., 2010), which allows them to have

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Fig. 4. A. Ordination triplot (RDA) of dominant, B. Abundant, and C. Rare species of urban landscape with association to

the most significant land cover and landscape indexes of Tuxtla Gutiérrez, total variance explained by the first two axes was

greater than 60 % and p < 0.001, for all cases.

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an advantage over other species in the use of

resources, avoid disturbances and improve their

communication (Sol et al., 2013).

Isolation and patch size of different land

covers interact to promote bird diversity within

a complex landscape mosaic to generate a bird

composition with a metacommunity structure.

The differential association of each of the sub-

sets of birds with landscape properties such as

the extent of tree cover, the heterogeneity of

covers and the intermixing of covers, allows

the urban landscape of TG to function as a

stepping stone of corridors facilitating disper-

sal and being a regulatory mechanism as an

environmental filter for species (Beninde et al.,

2015; Eyster et al. 2022), favoring their presence

in landscapes dominated by urban coverage

(Martin & Bonier, 2018).

The intense transformation and urbaniza-

tion of the Neotropical region, especially of

the tropical dry forest, is of important interest

for the conservation of biodiversity (Gordon

& Ornelas, 2000). The persistence of native

birds in cities and their conservation requires

a broad spatial scale perspective that allows us

to understand diversity patterns in the urban

landscape (McDonnell & Hahs, 2008). For the

city of TG, the remaining and sparse habitat

fragments of tree cover represent an important

refuge for native species, because several of

these birds are endemic or specialists in the

tropical dry forest habitat (Gordon & Ornelas,

2000). Finally, understanding the organization

patterns of species in a spatial context of trans-

formed landscapes allows us to recognize meta-

community structures that can help us detect

possible species association mechanisms and

environmental factors involved in the mainte-

nance of diversity in highly fragmented envi-

ronments such as tropical dry forests.

Ethical statement: The author(s) declared

no potential conflicts of interest with respect

to the research, authorship, and/or publica-

tion of this article, and followed all pertinent

ethical or legal procedures and requirements. A

signed document has been filed in the journal

archives.

ACKNOWLEDGMENTS

We would like to thank Universidad de

Ciencias y Artes de Chiapas for facilities in this

research, to the Programa para el Desarrollo

Profesional Docente (PRODEP) for a student

grant (AFG-AIC UNICACH_PTC-076) and

two anonymous reviewers for helpful com-

ments on versions of the manuscript.

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