Available online at www.sciencedirect.

com Landscape and Urban Planning 90 (2009) 189-195

This is the author’s version of a work accepted for publication by Elsevier in Landscape and Urban Planning (2009: volume 90,
issues 3-4, pages 189-195, DOI:10.1016/j.landurbplan.2008.11.003). The use of this document is for personal or academic use
only. © 2009 Elsevier B.V. All rights reserved

LIVING IN THE BIG CITY: EFFECTS OF URBAN LAND-USE ON

BIRD COMMUNITY STRUCTURE, DIVERSITY, AND COMPOSITION

Rubén Ortega-Álvarez1, Ian MacGregor-Fors2,*

Facultad de Ciencias, Universidad Nacional Autónoma de México, Ciudad Universitaria, Circuito exterior S/N,
1

México, D.F. 04510, México.

2
Laboratorio de Ecología Funcional, Centro de Investigaciones en Ecosistemas, Universidad Nacional Autónoma
de México, Campus Morelia, Antigua Carretera a Pátzcuaro 8701, Morelia 58190, Michoacán, México.
*
Author for correspondence: ian@oikos.unam.mx

Cities represent an important threat to biodiversity at different scales. Nevertheless, little is known on the
processes underlying such effects. In this paper we describe bird diversity, structure, and composition pat-
terns in different urban land-use categories. For this, we surveyed resident bird communities in four rep-
resentative land-use categories of southwestern Mexico City. Our results show that bird communities vary
greatly along the different studied urban land-uses, which represent an urbanization development gradient.
Bird communities were highly dominated by few generalist species in areas with commercial components,
while showed to have higher evenness values in green areas. Bird species richness decreased and bird abun-
dances increased with urbanization intensity. Also, our results indicate that bird species richness and abun-
dance values are sensible to site-specific habitat characteristics. Although we did not find a clear pattern of
taxonomic homogenization related to urbanization, our results show that urbanization development entails
the functional homogenization of bird communities. Thus, based on our results, we suggest three urban
planning and management activities: (1) regulate land-use change related to urbanization; (2) increase the
number of green areas within the city; (3) establish bird monitoring programs to identify focal areas that
need management and assist with ecological data for urban planning.
Keywords: Urban ecology, Land-use, Urban planning, Biodiversity, Neotropical bird communities.

1. INTRODUCTION

Urbanization replaces native habitats with new man-
made systems where natural and anthropogenic com-
ponents interact (Pickett et al., 1997). Such novel so-
cio-ecosystems encompass perdurable infrastructure,
and therefore the disturbance generated by the settle-
ment of urban areas is long-term and hardly reversible
(McKinney, 2002). Thus, the replacement of natural
habitats related to urbanization processes can lead to
species extinction and biotic homogenization (Czech
et al., 2000; Marzluff et al., 2001; McKinney, 2002;
Landscape and Urban Planning 90, 189-195
Olden et al., 2005), representing a threat to biodiversity
at different scales (Turner et al., 2004).
Urban systems comprise a great variety of compo-
nents, deriving in highly heterogeneous matrixes. Such
habitat variability is generated by: (1) the human activi-
ties carried out within an area; (2) its infrastructure; (3)
the vegetation components (Pickett et al., 1997). Bird
communities respond to this environmental variation
in several ways (Blair, 1996, 2001). In general, urban
bird communities include less species and higher abun-
dances than those from natural habitats (Gavareski,
1976; Beissinger and Osborne, 1982; Rosenberg et al.,
1987; Mills et al., 1989; Jokimäki and Suhonen, 1998).
Yet, little is known on the processes underlying such
patterns (Shochat et al., 2006). Generating information
that allows understanding the effects of urbanization
on the physical and biological components of ecosys-
tems could assist on the comprehension of the factors
determining avian presence and abundance within cit-
ies (Jokimäki and Suhonen, 1998; Miller et al., 2001).
In this study we evaluated the effect that various
urban land-use categories have on resident bird com-
munities, focusing on their diversity values, structure,
and composition. In particular, we sampled four repre-
sentative land-use categories in southwestern Mexico
City Metropolitan Area (referred as Mexico City here-
after), where we focused on the relationship between
bird community values and vegetation, urban structure,
predatory pressure, and human activity. We expected
that if environmental, resource, and human attributes
are substantially different among urban land-uses, the
diversity, structure, and/or composition of bird com-
munities should also vary accordingly.
2. MATERIALS AND METHODS
2.1. Study area
Mexico City is one of the largest metropolitan areas in
the world. At present, the city covers an approximate
area of 1800 km2 and houses 18,494,381 people (INE-
GI, 2006). Our study was conducted in the southwest-
ern section of the city, where the rough topography fa-
vored the establishment of diverse ecosystems before
urbanization (Rzedowski and Rzedowski, 2005). We se-
lected the southwestern section of the city to carry out
this study due to its high vertebrate diversity (Flores-
Villela and Gerez, 1994). Unfortunately, the biodiver-
sity encompassed within the city is constantly threat-
ened by the dynamics of population growth as a result
of its geographic expansion (Rojas, 2004). Although
the area where Mexico City was settled has changed
drastically since pre-Columbian times, it still remains
highly heterogeneous, but predominantly urban (Agui-
lar, 2004). Thus, similar to other Latin–American cit-
ies, Mexico City is basically categories: (1) commercial;
(2) residential; (3) industrial; (4) green areas.
2.2. Survey site characterization
To characterize sampling points, we measured urban
Landscape and Urban Planning 90, 189-195
structure, vegetation, predatory pressure, and human
activity in a 25m radius area. In particular, vegetation
structure was evaluated by: (1) number of vegetation
strata (tree, shrub, herbaceous plants); (2) tree spe-
cies richness; (3) tree abundance; (4) tree diameter at
breast height (DBH); (5) tree cover; (6) tree height;
(7) shrub species richness; (8) shrub cover; (9) shrub
height; (10) herbaceous plant cover; (11) herbaceous
plant height. Urban structure, human activity, and
potential human-related predatory pressure were de-
scribed by: (1) building height; (2) built cover; (3) pass-
ing cars/min; (4) pedestrians/min; (5) abundance of
dogs and cats.
2.3. Bird surveys
Resident landbirds were sampled during summer of
2007 from 07:00 to 10:00 in southwestern Mexico
City. For bird surveys, we conducted 10 min unlim-
ited radius point counts (following Ralph et al., 1993,
1995). Point counts were located in a minimum dis-
tance of 250m from each other in order to assure data
independence (Ralph et al., 1993; Huff et al., 2000).
We sampled 160 point counts in four urban land-use
categories: 40 in commercial areas, 40 in residential–
commercial areas, 40 in residential areas, and 40 in
green areas. Because industrial areas in Mexico City
are mainly located in its northern section, we did not
include this urban land-use category in our study. To
differentiate land-uses accurately, we first discriminat-
ed urbanized from non-urbanized areas (parks) using
the classification proposed by Marzluff et al. (2001).
Then, to distinguish between commercial, residential–
commercial, and residential land-uses, we used the
presence/absence of commercial components.
2.4. Data analysis
To assure that our survey effort was enough to record
a representative sample of southwestern Mexico City
bird communities, we computed two species predic-
tion analyses (SPADE; Chao and Shen, 2006). First,
we calculated the number of predicted species using
an abundance-based coverage estimator (ACE; Chao
and Shen, 2006). Second, we estimated the number of
new predicted species in a further survey using a mul-
tinomial model (cut-off point = 10), with a sample
size equal to five surveys.
We assessed differences in bird community struc-
ture by comparing the steepness of bird community
evenness/dominance slopes, using a species rank/
abundance plot approach (as suggested by Magurran,
2003). To evaluate statistical differences of the even-
ness/dominance slopes, we performed ANCOVA and
Newman–Keuls post hoc range tests. Because bird
abundance values widely differed between the studied
urban land-uses, values were transformed (log10).
In order to compare bird richness values between
the studied urban land-uses, we computed their spe-
cies richness statistical expectation (Sobs Mao Tau
±95% CI) using EstimateS (Colwell, 2005). This ex-
pectation is calculated from the repeated re-sampling
of all polled samples, and therefore allows statistical
comparisons between treatments/habitats (Gotelli
and Colwell, 2001). To test if urban land-uses affected
the abundance of birds, we used ANOVA. Bird abun-
dance values were transformed (log10) to fit a normal
distribution. We assessed species turnover rates among
urban land-uses with the Jaccard similarity coefficient
(Jaccard, 1912). Although several species turnover
indexes have also been proposed to measure species
turnover rates, the Jaccard similarity coefficient is rec-
ommended for evaluating species turnover rates when
comparisons follow a continuous approach (Koleff et
al., 2003).
We classified birds in trophic groups. For this, we
used two parameters: (1) primary feeding resource; (2)
body size (length). We used the variable ‘body size’ to
separate the wide ecological and functional roles ex-
isting between species from a same ‘primary feeding
resource group’. Primary feeding resource and body
size were determined bibliographically (Peterson Mul-
timedia Guides, 1996; Bull and Farrand, 1997). We per-
formed a multivariate cluster analysis to determine size
groups for each primary feeding resource (referred
as ‘trophic groups’ hereafter), using a 95% similarity
fringe to separate clusters. For comparing the trophic
composition of bird communities from the studied
urban land-uses, we conducted an abundance-based
Bray–Curtis multivariate cluster analysis (Bray and Cur-
tis, 1957), using BioDiversity Professional (McAleece,
1997).
To assess the effect that habitat attributes have
on bird species richness and abundances, we com-
puted stepwise multiple regressions. We seek for pos-
sible correlations between variables. When two or
more variables showed significant strong correlations
(R>0.5, P<0.05), we excluded those variables with low
variance and/or little significance for birds from the
analysis. Because two urban-exploiter species, the Rock
Landscape and Urban Planning 90, 189-195
Pigeon (Columba livia) and the House Sparrow (Passer
domesticus), comprised nearly one half of the total bird
abundance recorded in this study, we excluded their
abundances in another multiple regression analysis to
evaluate their effect. Thus, we performed three mul-
tiple regressions analyses for bird abundance, one with
the total recorded bird abundances, one with native
bird abundances, and one with Rock Pigeons-House
Sparrows.
3. RESULTS
3.1. Bird surveys
Results from the species prediction analyses show that
our survey effort was enough to record a representa-
tive sample of the bird communities from southwest-
ern Mexico City. The ACE revealed that our surveys
recorded 91% of the average predicted species. Rein-
forcing this result, the estimated number of new spe-
cies in a further survey was <1 species in all cases (data
not shown). We recorded 58 resident landbird species
of six trophic groups: insectivore (46.6%), granivore
(22.4%), omnivore (13.8%), nectarivore (8.6%), frugi-
vore (6.9%), and carnivore (1.7%). Of them, three are
considered as endangered by the Mexican government
(SEMARNAT, 2002), eight are endemic, and other
eight are quasi-endemic to Mexico (Howell and Webb,
1995; Sibley, 2001) (Table 1).
3.2. Community structure
The structure of bird communities recorded at the
studied urban land-uses differed. The slopes from
the species rank/abundance plots showed to be statis-
tically different among the studied urban land-use cat-
egories (ANCOVA: F3,131=15.36, P<0.001), showing
higher evenness in less developed urban land-uses (Fig.
1). Such differences were only significant when com-
paring green and commercial areas (Newman–Keuls:
P=0.034). Not finding differences between the species
rank/abundance slopes from residential and residen-
tial–commercial areas with the other urban land-uses
suggest that residential and residential–commercial
areas represent the middle-section of an urbanization
intensity gradient that progressively affects the steep-
ness of the evenness/dominance slopes of bird com-
munities.
3.3. Diversity values
Bird species richness decreased from green to com-
Table 1. Resident landbird species recorded in four urban land-use categories from southwestern Mexico
City. GA, green areas; RES, residential areas; R–C, residential–commercial areas; COM, commercial ar-
eas.

Family Species1,2,3 Urban land-uses

GA RES R-C COM
Odontophoridae Dendrortyx macroura (G)**P •
Columbidae Columba livia (O) • • •
Columbina inca (G) • • • •
Trochilidae Cynanthus latirostris (N)* • • •
Hylocharis leucotis (N) • • • •
Amazilia beryllina (N) • • • •
Lampornis clemenciae (N)* • • • •
Picidae Melanerpes formicivorus (O) • •
Picoides scalaris (I) •
Picoides villosus (I) •
Tyrannidae Contopus pertinax (I) •
Empidonax occidentalis (I)* • • •
Empidonax fulvifrons (I) •
Empidonax sp. (I) •
Tyrannus vociferans (I) •
Laniidae Lanius ludovicianus (C) •
Corvidae Cyanocitta stelleri (O) •
Aphelocoma ultramarina (O) • •
Hirundinidae Tachycineta thalassina (I) •
Hirundo rustica (I) • • • •
Paridae Poecile sclateri (I)* •
Aegithalidae Psaltriparus minimus (I) • • • •
Sittidae Sitta carolinensis (I) •
Sitta pygmaea (I) •
Certhiidae Certhia americana (I) •
Troglodytidae Thryomanes bewickii (I) • • • •
Troglodytes aedon (I) • •
Regulidae Regulus satrapa (I) •
Turdidae Myadestes occidentalis (I)P •
Catharus occidentalis (I)** •
Turdus assimilis (F) •
Turdus rufopalliatus (F)* • • • •
Turdus migratorius (F) • • • •
Mimidae Toxostoma curvirostre (I) • • •
Melanotis caerulescens (I)**P • •
Ptilogonatidae Ptilogonys cinereus (F)* • • •
Peucedramidae Peucedramus taeniatus (I) •
Parulidae Ergaticus ruber (I)** •
Myioborus miniatus (I) • •
Basileuterus rufifrons (I)* •
Basileuterus belli (I) •
Emberizidae Diglossa baritula (N) • • • •
Atlapetes pileatus (G)** •
Landscape and Urban Planning 90, 189-195
 Table 1. Continued.

Family Species1,2,3 Urban land-uses

GA RES R-C COM
Buarremon virenticeps (G)** •
Pipilo maculatus (G) • •
Pipilo fuscus (O) • • •
Oriturus superciliosus (G)** • • •
Melospiza melodía (G) • • • •
Junco phaeonotus (G)* • •
Cardinalidae Pheucticus melanocephalus (G) • • • •
Icteridae Quiscalus mexicanus (O) • • •
Molothrus aeneus (O) • • • •
Icterus abeillei (I)** • • • •
Fringillidae Carpodacus mexicanus (G) • • • •
Loxia curvirostra (G) •
Carduelis pinus (G) • •
Carduelis psaltria (G) • • • •
Passeridae Passer domesticus (O) • • • •

a
Trophic group: (I)=insectivore; (G)=granivore; (O)=omnivore; (F)=frugivore;
(N)=nectarivore; (C)=carnivore.
b
Endemism: *=quasi-endemic to Mexico; **=endemic to Mexico.
c
Status of endangerment sensu SEMARNAT (2002): P=under special protection.

mercial areas. When we compared the species rich- eas with commercial elements are highly similar. When
ness statistical expectations using a cut-off point of all studied land-uses were compared to green areas,
549 individuals, bird species richness recorded in residential areas showed the closest taxonomic simi-
green areas (55±4.74 species) was significantly higher larity, followed by commercial and residential–com-
when compared to those from the other studied urban mercial areas. Thus, the taxonomic similarity of the re-
land-uses. The statistical expectation of bird species corded bird communities, measured using the Jaccard
richness from residential areas (31.49±4.30 species) similarity coefficient, showed that the studied urban
only showed statistical differences with the ones from land-uses represent an urban development gradient
residential–commercial (22.78±3.58 species) and com- that molds the composition of bird communities in
mercial areas (20.22±4.17 species), which did not show southwestern Mexico City (Table 2). These results are
significant differences between them (Fig. 2). similar to those obtained from the Bray–Curtis multi-
Bird abundances increased significantly from areas variate cluster analysis, which showed that commercial
with low to high urbanization development (ANOVA: and residential–commercial areas are highly similar.
F3,156=2.67, P=0.049). The highest bird abundance On the other hand, residential areas showed higher
value was recorded in commercial areas (26.70±5.68 similarity to green areas than to commercial and resi-
individuals), followed by residential–commercial areas dential–commercial areas (Fig. 3).
(19.95±1.18 individuals), residential areas (17.22±0.92
individuals), and green areas (15.05±1.90 individuals; 3.5.Trophic composition
Fig. 2). We recorded bird species of six avian trophic groups
in this study, including frugivores, granivores, insecti-
3.4. Taxonomic composition vores, nectarivores, omnivores, and carnivores (repre-
Species turnover rates assessed using the Jaccard simi- sented by one single species).We differentiated the car-
larity coefficient show that bird communities from ar- nivore species from insectivores due to the ecological

Landscape and Urban Planning 90, 189-195


Figure 1. Species rank/abundance plots for the studied urban
land-uses. Comparisons of the evenness/dominance slopes only
showed differences between green and commercial areas, suggest-
ing that residential and residential–commercial areas comprise the
middle-section of an urbanization intensity gradient. Thus, our
results suggest that such gradient affects bird community even-
ness/dominance progressively.
differences between ‘strict insectivores’, such as war-
blers or tyrant-flycatchers, and the recorded carnivore
bird species (Loggerhead Shrike—Lanius ludovicianus),
which basically feeds on small vertebrates and large in-
vertebrates (Percevia Field Guides, 2007).
The abundance of species pertaining to the estab-
lished trophic groups differed between the studied ur-
ban land-uses (Table 3). While omnivores dominated
bird communities in commercial, residential–commer-
cial and residential areas, the most abundant trophic
groups in green areas were granivores and insectivores.
The multivariate cluster analysis revealed that the tro-
phic composition of bird communities was essentially
identical in commercial, residential–commercial and
residential areas, while showed slight differences in re-
lation to green areas (Fig. 3).
3.6. Relationships with habitat attributes
Habitat attributes differed between the studied urban
land-uses (Table 4). Multiple regression analyses re-
vealed that bird species richness values in southwest-
ern Mexico City are significantly and positively related
to shrub cover, shrub height, and herbaceous plant
height, and negatively related to passing cars/min (Ta-
ble 5). On the other hand, total bird abundance was
Landscape and Urban Planning 90, 189-195
Figure 2. Bird diversity values. Bird species richness was higher
in green areas, while bird abundance was higher in developed
land-uses.GA: green areas; RES: residential areas; R–C: residen-
tial–commercial areas; COM: commercial areas. Segmented lines
are displayed in order to represent the pattern of both variables
throughout the urban land-use gradient.
positively related to pedestrians/min, and negatively
related to passing cars/min (Table 6(a)). The abun-
dance of native bird species showed to be positively
related to shrub height and tree species richness, while
showed negative relationship to passing cars/min (Ta-
ble 6(c)). Finally, the abundance of both exotic bird
species, the Rock Pigeon and the House Sparrow, was
positively related to pedestrians/min, and negatively
related to tree abundance and passing cars/min (Table
6(b)).
4. DISCUSSION AND CONCLUSIONS
Our results provide evidence that the studied urban
land-use categories of southwestern Mexico City af-
fect the bird communities they shelter. Bird communi-
ties showed to be highly dominated by a few urban-
exploiter species in areas that comprise commercial
components, while were fairly even in green areas.
This pattern was supported by the recorded bird com-
munity diversity values, which showed decreasing bird
species richness and increasing bird abundance in re-
lation to the level of urban development. Our results
suggest that these patterns are, in part, explained by
habitat attributes. Such differences in the urban habi-
tats were reflected by the functional composition of
their bird communities, being identical in residential,
commerial, and residential–commercial areas. In this
section we first discuss bird community structure pat
 Table 2. Bird taxonomic similarity in different urban land-uses. The Jaccard similarity coefficient showed
that bird communities from commercial and residential–commercial areas are highly similar, while those
from green areas are highly dissimilar to the other studied land-uses. On the other hand, bird communi-
ties from residential areas ere more similar to those from green areas than in relation to the ones from
commercial and residential–commercial areas.

Jaccard similarity coefficient (%)

Green area Residential Residential–commercial
Green area –
Residential 58.09 –
Residential–commercial 38.57 68.46 –
Commercial 36.52 57.48 71.27

Table 3. Trophic group abundances. Values represent the sum of all recorded landbird individuals from
each trophic group.

Size categorya Bird abundances in land-use categories

Green area Residential Residential–commercial Commercial

Carnivore 4 0 0 0
Frugivore1 3 15 4 0
Frugivore2 43 51 27 31
Granivore1 95 155 126 96
Granivore2 46 23 19 19
Granivore3 36 32 126 67
Granivore4 4 0 0 0
Insectivore1 8 0 0 0
Insectivore2 124 47 19 41
Insectivore3 24 20 37 30
Insectivore4 7 13 3 2
Nectarivore1 50 52 31 22
Nectarivore2 7 3 2 1
Omnivore1 58 218 335 346
Omnivore2 30 43 11 28
Omnivore3 16 17 56 375
Omnivore4 47 0 2 10

Numbers displayed after trophic guilds represent the size category resultant of the
a

multivariate cluster analysis (ascendant order).

terns in the studied urban land-use categories. Second- Bird communities recorded in southwestern Mex-
ly, we focus on the patterns found for bird diversity ico City showed to be highly dominated by few ur-
values and differences in species composition. Thirdly, ban-exploiter bird species in highly developed areas,
we analyze the relationships found between habitat at- while were fairly even in green areas. This result of-
tributes and bird diversity values. Finally, we suggest fers an insight on the distribution of the resources
three urban management and planning actions that and conditions offered to birds in each of the stud-
could help on the maintenance and promotion of ied urban land-uses, suggesting that green areas offer
complex bird communities in urban ecosystems. a wider spectrum of resources available for granivore
Landscape and Urban Planning 90, 189-195

Figure 3. Bird community similarity in different urban land-uses.
(a) The taxonomic approach shows that commercial and residen-
tial–commercial areas are highly similar, while residential areas
showed higher similarity to green areas. These results are similar
to those obtained using the Jaccard similarity coefficient. (b) The
functional approach showed that commercial, residential–com-
mercial, and residential areas comprise functionally identical bird
communities, which differed from those recorded in green areas.
Numbers aside each node represent its similarity value (%). GA:
green areas; RES: residential areas; R–C: residential–commercial
areas; COM: commercial areas.
and insectivore species (as recorded in a Mexican sub-
urb; MacGregor-Fors, 2008). On the other hand, the
dominance of omnivore and granivore species in resi-
dential, residential–commercial, and commercial land-
uses, demonstrates that these urban land-uses tend to
select for generalist species, as ecorded for other bird
communities in the US, Canada, and Europe (Emlen,
1974; Beissinger and Osborne, 1982; Rosenberg et al.,
1987; Clergeau et al., 1998; Jokimäki and Kaisanlahti-
Jokimäki, 2003).
Bird community structure results were supported
by the analysis of bird diversity. Our results show that
bird species richness differs in the different urban
land-uses of southwestern Mexico City. Similar to the
results of other urban ecology studies (e.g., Emlen,
1974; Beissinger and Osborne, 1982; Nocedal, 1987;
Clergeau et al., 1998; McKinney, 2002; Melles et al.,
2003), we found that bird species richness decreases
Landscape and Urban Planning 90, 189-195
with the intensity of urban disturbance. This pat-
tern can be mainly explained by three factors. First,
urban systems enclose great environmental variation
(Sukopp, 1998), being its vegetation component cru-
cial for birds (Emlen, 1974; Gavareski, 1976; Mills et
al., 1989; MacGregor-Fors, 2008). Second, some na-
tive bird species are incapable of invading urban areas
successfully due to the lack of resources and/or suit-
able habitat conditions, reason why they are generally
absent in highly developed sites (Blair, 2004; Clergeau
et al., 2006). Third, highly urbanized sites, represented
in this study by commercial areas, are locations where
pollution and anthropogenic disturbances are intense.
Thus, highly developed areas represent a threat to
those species that are sensitive to such stressor agents
(McKinney, 2002). Contrary to our bird species rich-
ness results, bird abundances increased with urbaniza-
tion intensity. This pattern is also similar to those re-
ported for other bird communities around the world
(e.g., Emlen, 1974; Beissinger and Osborne, 1982;
Mills et al., 1989; Clergeau et al., 1998; Shochat, 2004;
Chace andWalsh, 2006). Such an increase is closely re-
lated to the demographic growth of urban exploiters,
which tend to replace urban-adaptable bird species
and drive urban bird communities to a homogenized
state (McKinney, 2006). Thus, because highly devel-
oped urban areas are associated with high food re-
source predictability (Shochat, 2004), urban landuses
with commercial elements represent ideal habitats for
the demographic explosion of those species able to
exploit the abundant food resources associated to hu-
man litter.
Species turnover rates between the studied urban
land-uses were strongly related to urban habitat char-
acteristics. Our results show that urban green areas
comprise high vegetation complexity, followed by
residential areas. These results clearly show that bird
communities do respond to habitat attributes within
urban areas, and therefore can be used as indicators
of the ecological complexity of specific urban lo-
calities. Surprisingly, avian trophic groups responded
differently to the studied land-uses. In this study, ur-
banization represented a functional threshold for bird
communities, where regardless of the degree of ur-
banization disturbance, outside green areas bird com-
munities were functionally homogeneous. While green
areas comprised functionally complex bird communi-
ties, developed urban land-uses exhibited functionally
equivalent ones. We believe that this pattern is related
 Table 4. Habitat similarity between the studied urban land-uses. The multivariate cluster analysis showed
that residential and residential–commercial areas are highly similar, while the highest dissimilarity was
found between green areas and commercial areas.

Similarity (%)
Green area Residential Residential–commercial
Green area –
Residential 73.95 –
Residential–commercial 38.57 71.34 89.45 –
Commercial 57.00 73.02 77.16

Table 5. Relationship between habitat attributes and bird species richness.

General model (R = 0.57; F4,153 = 18.82; P < 0.001)

Beta S.E. t(153) P

Intercept 4.89420 0.000002

Shrub cover 0.22519 0.07804 2.88544 0.004473
Shrub height 0.17303 0.07324 2.36242 0.019413
Herbaceous height 0.19003 0.07647 2.48486 0.014036
Passing cars/min −0.26537 0.07103 −3.73580 0.000263

to two main factors. First, specific food resources of-
fered to birds in urban green areas, summed to the
complexity, quality, and abundance of nesting, roost-
ing, and feeding sites can maintain complex bird com-
munities. Second, urban green areas comprise areas
where human disturbance is lower than in other urban
land-uses, variable that has been positively related to
avian functional homogenization in fragmented land-
scapes (Devictor et al., 2008).
Species richness values recorded in our study area
were significantly related to three vegetation attributes:
(1) shrub cover; (2) shrub height; (3) herbaceous plant
height. This result is not surprising, since the vegeta-
tion component has been positively related to bird spe-
cies richness in the past (Gavareski, 1976; Melles et al.,
2003; MacGregor-Fors, 2008). Particularly, shrubs of-
fer various benefits to birds, such as escape cover, nest-
ing sites, and foraging resources (Savard et al., 2000;
Fernández-Juricic et al., 2001; Melles et al., 2003). On
the other hand, areas with high herbaceous plants
within Mexico City comprise non-managed lots that
recall natural shrublands or grasslands, benefiting gra-
nivores and other bird species that nest on the ground
(Gering and Blair, 1999; Jokimäki and Huhta, 2000;
MacGregor-Fors, 2008). In contrast, species richness
showed to be negatively related to human activity. This
relationship is noticeable, since car activity generates
noise, collisions, and/or air pollution, which can drive
urban-adaptable birds away from areas with heavy traf-
fic (see Chace and Walsh, 2006 and references there-
in).
Total bird abundance, greatly influenced by the
abundance of House Sparrows and Rock Pigeons, ex-
hibited strong relationships with human activity. Con-
sequently, when native bird abundance was analyzed,
relationships were quite different to those from the
total species richness analysis, showing positive rela-
tionships with vegetation components, and negative
relationships with urbanization intensity related vari-
ables. These results show that the abundance of the
few urban-exploiter species that dominate urban bird
communities can bias analyses when considering total
urban bird abundance. Thus, special care needs tobe
taken when including the abundance of urban-exploit-
er species in ecological analyses.
Urbanization has been a force of land transforma-
tion for centuries. Although human kind will probably
never get to undo the negative effects that urbaniza-
tion has had on biodiversity, urban management and
planning activities based on urban ecology knowledge
could mitigate such effects (Jokimäki, 1999; Turner et
al., 2004). The present study was only carried out in

Landscape and Urban Planning 90, 189-195

the southwestern section of Mexico City. However,
results included in this paper are robust and confirm
that some patterns reported previously by others in
the US, Canada, and Europe also apply in one of the
largest metropolis of the world. Although some urban
ecology patterns are consistent among studies, efforts
to achieve urban planning and management recom-
mendations are not consistent among countries. Spe-
cifically, Mexican cities grow following inadequate ur-
ban planning (e.g., Morelia; López et al., 2001), where
the establishment of new urban green areas appears
not to be a priority. Thus, although urban planning in
developed countries do contemplate the importance
of urban-based land-use change and the establish-
ment of urban green areas, based on our results, we
suggest that Mexican urban planning policies should
focus on: (1) regulating land-use change within urban
areas following a framework of environmental sustain-
ability; (2) increasing the quantity and complexity of
the vegetation cover in residential gardens, sidewalks,
and median strips with the aim of generating urban
green networks. Also, because each city is a unique sys-
tem, its management and planning activities should be
continuously evaluated to measure their effectiveness.
Thus, we recommend monitoring the environmental
quality of cities using birds as bio-indicators. Such
monitoring programs should use sensitive sampling
methods, such as point-counts (following Ralph et al.,
1993), and would be highly efficient if were included
in a volunteer-based project, such as the Tucson Bird
Count (Turner, 2003). These urban management and
planning activities could assist on the maintenance and
promotion of native bird species richness in urban ar-
eas, as well as increasing the environmental quality of
the urban systems we live in.
ACKNOWLEDGEMENTS
This study was partially granted by the project Manejo
de Ecosistemas y Desarrollo Humano en la Cuenca
del Río Magdalena, Universidad Nacional Autónoma
de México. We thank Erick de la Barrera for review-
ing our manuscript, Ana Lagunes-Gasca for help with
field-work, and two anonymous reviewers that im-
proved the quality of our manuscript.
LITERATURE CITED
Aguilar, A. 2004. Procesos metropolitanos y grandes ciudades:
Dinámicas recientes en México y otros países. H. Cámara de
Diputados, Universidad Nacional Autónoma de México, Con-
sejo Nacional de Ciencia y Tecnología, Miguel Ángel Porrúa,
Landscape and Urban Planning 90, 189-195
México, D.F. p. 536.
Beissinger, S.R., Osborne, D.R., 1982. Effects of urbanization on
avian community organization. Condor 84, 75–83.
Blair, R.B., 1996. Land use and avian species diversity along an ur-
ban gradient. Ecological Applications 6, 506–519.
Blair, R.B., 2001. Birds and butterflies along urban gradients in two
ecoregions of the United States: is urbanization creating a ho-
mogeneous fauna? In: Marzluff, J.M., Bowman, R., Donnely,
R. (Eds.), Avian Conservation and Ecology in an Urbanizing
World. Kluwer Academic, Boston, pp. 33–56.
Blair, R.B., 2004. The effects of urban sprawl on birds at multiple
levels of biological organization. Ecology and Society 9, 2 (on-
line).
Bray, J.R., Curtis, J.T., 1957. An ordination of the upland forest
communities of Southern Wisconsin. Ecological Monographs
27, 325–349.
Bull, J., Farrand, J., 1997. National Audubon Society Field Guide
to North American Birds—Eastern Region. Alfred A. Knopf,
New York, p. 800.
Chace, J.F., Walsh, J.J., 2006. Urban effects on native avifauna: a
review. Landscape and Urban Planning 74, 46–69.
Chao, A., Shen, T.J., 2006. Program SPADE (species prediction and
diversity estimation). Version 3.1. http://chao.stat.nthu.tw.
Clergeau, P., Savard, J.P.L., Mennechez, G., Falardeau, G., 1998.
Bird abundance and diversity along an urban-rural gradient: a
comparative study between two cities on different continents.
Condor 100, 413–425.
Clergeau, P., Croci, S., Jokimäki, J., Kaisanlahti-Jokimäki, M.L.,
Dinetti, M., 2006. Avifauna homogenisation by urbanisation:
analysis at different European latitudes. Biological Conserva-
tion 127, 336–344.
Colwell, R.K. 2005. EstimateS: statistical estimation of species rich-
ness and shared species from samples. Version 7.5. http://purl.
oclc.org/estimates.
Czech, B., Krausman, P.R., Devers, P.K., 2000. Economic associa-
tions among causes of species endangerment in the United
States. BioScience 50, 593–601.
R. Ortega-Álvarez, I. MacGregor-Fors / Landscape and Urban
Planning 90 (2009) 189–195 195
Devictor, V., Julliard, R., Clavel, J., Jiguet, F., Lee, A., Couvet, D.,
2008. Functional biotic homogenization of bird communities
in disturbed landscapes. Global Ecology and Biogeography 17,
252–261.
Emlen, J.T., 1974. An urban bird community in Tucson, Arizona:
derivation, structure, regulation. Condor 76, 184–197.
Fernández-Juricic, E., Jimenez, M.D., Lucas, E., 2001. Bird toler-
ance to human disturbance in urban parks of Madrid (Spain):
Management implications. In: Marzluff, J.M., Bowman, R.,
Donnely, R. (Eds.), Avian Conservation and Ecology in anUr-
banizingWorld. Kluwer Academic, Boston, MA, pp. 259–273.
Flores-Villela, O., Gerez, P., 1994. Biodiversidad y conservación en
México: Vertebrados, vegetación y uso de suelo. Ediciones Téc-
nico Científicas, México, D.F. p. 463.
Gavareski, C.A., 1976. Relation of park size and vegetation to urban
bird populations in Seattle,Washington. Condor 78, 375–382.
Gering, J.C., Blair, R.B., 1999. Predation on artificial bird nests
along an urban gradient: predatory risk or relaxation in urban
environments? Ecography 22, 532–541.
Gotelli, N.J., Colwell, R.K., 2001. Quantifying biodiversity: proce-
dures and pitfalls in the measurement and comparison of spe-
cies richness. Ecology Letters 4, 379–391.
Howell, S.N.G.,Webb, S., 1995. A Guide to the Birds of Mexico and
 Northern Central America. Oxford University Press, Oxford,
p. 1010.
Huff, M.H., Bettinger, K.A., Ferguson, M.L., Brown, M.J., Altman,
B., 2000. A Habitatbased Point-count Protocol for Terrestrial
Birds, Emphasizing Washington and Oregon. U.S. Department
of Agriculture, Portland, p. 39.
(INEGI) Instituto Nacional de Estadística, Geografía e Informática,
2006. Cuaderno Estadístico de la Zona Metropolitana del Valle
de México. www.inegi.gob.mx.
Jaccard, P., 1912. The distribution of the flora in the alpine zone.
New Phythologist 11, 37–50.
Jokimäki, J., Suhonen, J., 1998. Distribution and habitat selection of
wintering birds in urban environments. Landscape and Urban
Planning 39, 253–263.
Jokimäki, J., 1999. Occurrence of breeding bird species in urban
parks: effects of park structure and broad-scale variables. Urban
Ecosystems 3, 21–34.
Jokimäki, J., Huhta, E., 2000. Artificial nest predation and abundance
of birds along an urban gradient. Condor 102, 838–847.
Jokimäki, J., Kaisanlahti-Jokimäki, M.L., 2003. Spatial similarity of
urban bird communities: a multiscale approach. Journal of Bio-
geography 30, 1183–1193.
Koleff, P., Gaston, K.J., Lennon, J.J., 2003. Measuring beta diver-
sity for presence–absence data. Journal of Animal Ecology 72,
367–382.
López, E., Bocco, G., Mendoza, M., Duhau, E., 2001. Predicting
land-cover and landuse change in the urban fringe: a case in Mo-
relia city, Mexico. Landscape and Urban Planning 55, 271–285.
MacGregor-Fors, I., 2008. Relation between habitat attributes and
bird richness in a western Mexico suburb. Landscape and Urban
Planning 84, 92–98.
Magurran, A.E., 2003. Measuring Biological Diversity. Blackwell
Publishing, Oxford, p. 260.
Marzluff, J.M., Bowman, R., Donnely, R., 2001. A historical perspec-
tive on urban bird research: trends, terms, and approaches. In:
Marzluff, J.M., Bowman, R., Donnely, R. (Eds.), Avian Conser-
vation and Ecology in an Urbanizing World. Kluwer Academic,
Boston, MA, pp. 1–17.
McAleece, N., 1997. BioDiversity Professional. Version 2. http://
www.sams.ac.uk/activities/downloads/downloads.htm.
McKinney, M.L., 2002. Urbanization, biodiversity and conservation.
BioScience 52, 883–890.
McKinney, M.L., 2006. Urbanization as a major cause of biotic ho-
mogenization. Biological Conservation 127, 247–260.
Melles, S., Glenn, S., Martin, K., 2003. Urban bird diversity and
landscape complexity: species–environment associations along a
multiscale habitat gradient. Conservation Ecology 7, 5 (online).
Miller, J.R., Fraterrigo, J.M., Hobbs, N.T., Theobald, D.N., Wiens,
J.A., 2001. Urbanization, avian communities, and landscape
ecology. In: Marzluff, J.M., Bowman, R., Donnely, R. (Eds.),
Avian Conservation and Ecology in an Urbanizing World. Klu-
wer Academic, Boston, MA, pp. 117–137.
Mills, G.S., Dunning, J.B., Bates, J.M., 1989. Effects of urbanization
on breeding bird community structure in Southwestern desert
habitats. Condor 91, 416–428.
Nocedal, J., 1987. Las comunidades de pájaros y su relación con la
urbanización en la Ciudad de México. In: Rapoport, E., López-
Moreno, I. (Eds.), Aportes a la Ecología Urbana de la Ciudad de
México. Editorial Limusa, México, D.F., pp. 73–109.
Olden, J.D., Douglas, M.E., Douglas, M.R., 2005. The human di-
mensions of biotic homogenization. Conservation Biology 19,
2036–2038.
Landscape and Urban Planning 90, 189-195
Percevia Field Guides, 2007. Field Guide to Birds of North Ameri-
ca. Percevia Field Guides. Available at http://identify.whatbird.
com/mwg/ /0/attrs.aspx.
Peterson Multimedia Guides, 1996. North American Birds (CD
ROM) Version 3.1. Houghton Mifflin Interactive, New York.
Pickett, S., Burch, W., Dalton, S., Foresman, T., Morgan, J., Rown-
tree, R., 1997. A conceptual framework for the study of human
ecosystems in urban areas. Urban Ecosystems 1, 185–199.
Ralph, C.J., Geupel, G.R., Pyle, P., Martin, T.E., DeSante, D.F., 1993.
Handbook of Field Methods for Monitoring Landbirds. U.S.
Department of Agriculture, Forest Service, Pacific Southwest
Research Station. Gen. Tech. Rep. PSW-GTR-144, Albany, CA.
p. 41.
Ralph, C.J., Droege, S., Sauer, J.R., 1995. Managing and Monitoring
Birds Using Point Counts: Standards and Applications. United
States Department of Agriculture, p. 169.
Rojas, E., 2004. Los desafíos de un continente urbano: La acción del
BID en desarrollo urbano.Washington, D.C., Banco de Desar-
rollo Interamericano, p. 60.
Rosenberg, K.V., Terrill, S.B., Rosenberg, G.H., 1987. Value of
suburban habitats to desert riparian birds. Wilson Bulletin 99,
642–654.
Rzedowski, G., Rzedowski, J., 2005. Flora fanerogámica del Valle de
México. Instituto de Ecología, A.C., Comisión Nacional para el
Conocimiento y Uso de la Biodiversidad, Pátzcuaro, Michoacán.
p. 403.
Savard, J.L., Clergeau, P., Mennechez, G., 2000. Biodiversity con-
cepts and urban ecosystems. Landscape and Urban Planning 48,
131–142.
(SEMARNAT) Secretaría de Medio Ambiente y Recursos Naturales,
2002. Norma Oficial Mexicana NOM-059-ECOL-2001, Protec-
ción Ambiental-Especies Nativas de México de Flora y Fauna
Silvestres-Categorías de Riesgo y Especificaciones para su Inclu-
sión, Exclusión o Cambio-Lista de Especies en Riesgo. Diario
Oficial de la Federación-Segunda Sección-Secretaría del Medio
Ambiente y Recursos Naturales, México, D.F. p. 171.
Shochat, E., 2004. Credit or debit? Resource input changes popula-
tion dynamics of city-slicker birds. Oikos 106, 622–626.
Shochat, E., Warren, P.S., Faeth, S.H., McIntyre, N.E., Hope, D.,
2006. From patterns to emerging processes in mechanistic ur-
ban ecology. Trends in Ecology and Evolution 21, 186–191.
Sibley, D.A., 2001. The Sibley Guide to Birds. Alfred A. Knopf, New
York, p. 544.
Sukopp, H., 1998. Urban ecology: scientific and practical aspects. In:
Breuste, J., Feldmann, H., Uhlmann, O. (Eds.), Urban Ecology.
Springer, Berlin, pp. 3–16.
Turner, W.R., 2003. Citywide biological monitoring as a tool for ecol-
ogy and conservation in urban landscapes: the case of the Tuc-
son Bird Count. Landscape and Urban Planning 65, 149–166.
Turner,W.R., Nakamura, T., Dinetti, M., 2004. Global urbaniza-
tion and the separation of human from nature. BioScience 54,
585–590.