Gonzalo A. Ramírez-Cruz’s research while affiliated with Arizona State University and other places

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Publications (19)


Unexpected spatial aggregation between two species of hummingbirds and their inconsistent spatial interactions with native and exotic plants in an urban ecological reserve
  • Article

March 2025

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23 Reads

Ibis

Gonzalo A. Ramírez‐Cruz

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J. Jaime Zúñiga‐Vega

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[...]

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Mariana Palencia‐Martínez

Hummingbirds play an important role as pollinators and are one of the primary examples of animal–plant coevolution. However, factors such as land‐use change, urbanization, exotic species introductions and the disappearance of native plants can negatively affect the persistence of hummingbird populations. Thus, understanding the ecological factors that may favour their presence is essential for their effective conservation in urban areas. We conducted repeated surveys in a native scrubland reserve within Mexico City from 2015 to 2018, and used conditional two‐species occupancy models to estimate a species interaction factor, and to analyse the effect of five native plants ( Agave salmiana , Echeveria gibbiflora , Opuntia lasiacantha , O. tomentosa and Wigandia urens ) and one exotic plant ( Leonotis nepetifolia ) on the occupancy probability of two hummingbirds: the Broad‐billed Hummingbird ( Cynanthus latirostris ) and the Berylline Hummingbird ( Saucerottia beryllina ). Considering that both of the hummingbird species could be competing for similar resources, we also tested the effect of the presence of the Berylline Hummingbird on the occupancy of the Broad‐billed Hummingbird, assuming its dominance based on larger body‐size. We found evidence of positive spatial interactions between plants and hummingbirds, but we did not find these to be consistent throughout the 3 years of monitoring, which could indicate a modification of the original hummingbird–plant interaction networks resulting from urban disturbance. In addition, there were no negative spatial interactions between the two hummingbirds. Instead, we detected an aggregation of both species during two separate seasons, probably owing to one species using the other as a cue for profitable sites. Even though the populations of these two hummingbird species can thrive in an urban ecosystem, changes in spatial interactions might reflect alterations on the structure and functioning of a community. Therefore, we consider it crucial to continue studying their population dynamics and their implications for the conservation of urban hummingbirds.



Keystone species in an urban environment: Do raptors control the Mexican gray squirrel in Mexico City?

September 2024

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50 Reads

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1 Citation

Acta Oecologica

Urbanization has forced animals to respond to novel environments. One of the basic questions in urban ecology is if ecological processes that occur in natural conditions also occur in urban environments. In natural ecosystems, key predators can limit prey abundance and have a widespread effect on trophic levels. Here, we examine a predator-prey interaction in Mexico City, an urban ecosystem where prey can be evaluated in the presence or absence of apex predators. Nearly all original ecosystems in Mexico City have been transformed for human purposes. Many of the green spaces in the city contain populations of the Mexican gray squirrel, a species native to this area. These green spaces are also inhabited by different species of raptors, the most frequent being Harris's hawk, Cooper's hawk, and sharp-shinned hawk. Little is known about the interactions between raptors and squirrels, particularly how different environmental factors influence this ecological relationship. We do know, however, that raptors prey on squirrels. We predicted that in parks where raptors were present, there would be fewer squirrels than in parks where there were no raptors. We studied the relationship between raptors and squirrels using occupancy models, which also allowed us to evaluate environmental factors that affect the presence of both squirrels and raptors. We also tested if the presence of raptors influenced the occupancy probability of squirrels, and vice versa. Lastly, we estimated the abundance of squirrels in parks, both where raptors were present and where they were absent. Contrary to our predictions, in our first two sampling periods we found a positive relationship between the presence of raptors and squirrel abundance, but this relationship was absent during our second two survey periods, which was due to a decline in squirrel abundance in parks where raptors were present. These results suggest that predator-prey interactions occur in urban settings much as they do under natural conditions, but that other factors unique to urbanization can also impact the abundance of raptors and squirrels. Our findings also suggest that raptors might be an effective means of controlling squirrel populations under some conditions.


Correction to: Dynamic occupancy analyses of native birds in an urban ecological reserve reveal seasonal changes in site occupancy and preference for adjacent urbanized areas
  • Article
  • Full-text available

July 2024

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13 Reads

Urban Ecosystems

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Location in Mexico of the urban ecological reserve where we conducted our study (Reserva Ecológica del Pedregal de San Ángel) including the surrounding urban areas. This reserve is divided into core conservation areas (colored in dark blue) and buffer areas (colored in green). All the surrounding areas represent gardens, buildings, parking lots, paved walking paths, and streets. Landmarks indicate the location of our 100 observation sites and are colored according to their location in either core, buffer, or urban areas. The number of observation sites within each type of area is shown within parentheses
Estimated relationships between initial occupancy probability (ψ) (i.e., occupancy during our first field season, which took place during May 2015) and some environmental variables for different species of native birds. Dashed gray lines and error bars denote 95% confidence intervals
Estimated relationships between colonization probability (𝛾) and some environmental variables for different species of native birds. Dashed gray lines and error bars denote 95% confidence intervals
Estimated relationships between local extinction probability (ε) and some environmental variables for different species of native birds. Dashed gray lines and error bars denote 95% confidence intervals
Model-averaged estimates of occupancy probabilities (ψ) for nine species of native birds and for all field seasons. Different symbols represent different climatic seasons. Error bars denote 95% confidence intervals
Dynamic occupancy analyses of native birds in an urban ecological reserve reveal seasonal changes in site occupancy and preference for adjacent urbanized areas

March 2024

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133 Reads

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3 Citations

Urban Ecosystems

Urban ecological reserves are large green areas immersed within cities where native and exotic species of plants and animals coexist. Here, we examined the environmental features that facilitate the occurrence of nine species of native birds in an urban ecological reserve located within one of the largest cities in the world, Mexico City. We also searched for changes in occupancy rates among the three distinct climatic seasons that occur in central Mexico: warm-dry, rainy, and cold-dry. Using data collected during four years and multi-season occupancy models, we found that most of our study species prefer the urbanized sites that surround the reserve over the core conservation areas. This urban affinity can be explained by the diverse vegetation that prevails in such urban sites, which offers a high habitat heterogeneity that facilitates the presence of bird species with distinct ecological needs. In contrast, the reserve consists of a relatively homogeneous xerophytic scrubland where a few species of shrubs and small trees are dominant. We also detected seasonal changes in five species, with highest occupancy during the warm-dry season of each year, which coincides with both their breeding season and the driest period of the year. This finding indicates that these birds find in the reserve and surrounding urban areas enough food and water during this limiting season as well as safe nesting sites. Our study provides evidence that some native birds can become urban exploiters and that the benefits that they obtain from urban settings are greatest during their breeding season.



Dynamic rates of site occupancy by native birds in an urban ecological reserve

September 2022

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63 Reads

Urban ecological reserves are relatively large green areas immersed within cities that serve the purpose of preserving the native species of plants and animals that still persist in urban settings. In this study, we examined temporal changes (among seasons and years) in site occupancy rates of nine species of native birds that inhabit an urban ecological reserve located within one of the largest cities in the world, Mexico City. Using data collected during 11 field seasons that spanned a four-year period and implementing multi-season occupancy models, we estimated occupancy, colonization, local extinction, and detection probabilities of all nine species. For more than half of our study species, the highest occupancy and colonization rates occurred at or near the urban areas that surround the reserve. Additionally, in three species local extinction probabilities were lowest also at these urban sites. This urban affinity can be explained by the diverse vegetation that prevails in the surrounding urban sites, which offers a high habitat heterogeneity that facilitates the presence of bird species with distinct ecological needs. In contrast, the ecological reserve consists of a relatively homogeneous xerophytic scrubland where a few species of shrubs and small trees are dominant. We did not detect long-term decreasing trends in site occupancy of any species. Instead, in three species we observed positive trends in occupancy during particular periods. Overall, our findings indicate that the combination of this urban ecological reserve with the green spaces of the surrounding urban areas harbors stable populations of native birds.


Predicted relationships between a shrub cover, b tree cover, and c distance to developed areas and colonization probabilities of the House Sparrow within the ecological reserve. Dotted lines represent confidence intervals (95%) of the estimations
Predicted relationships between distance to developed areas and extinction probabilities of the House Sparrow within the ecological reserve. Dotted lines represent confidence intervals (95%) of the estimations
Identifying management guidelines to control the invasive House Sparrow (Passer domesticus) within natural protected areas through the estimation of local colonization and extinction probabilities

December 2021

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84 Reads

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11 Citations

Biological Invasions

Invasive species hinder the conservation objectives of natural protected areas, particularly of those found within or nearby urban settlements. Identifying the habitat and landscape traits that determine the establishment and persistence of populations is essential for implementing effective management plans to control invasive species. We employed multi-season occupancy models to identify the habitat and landscape traits that determined the local colonization and extinction probabilities of an invasive bird (House Sparrow—Passer domesticus), in order to provide recommendations for controlling its population within a natural protected area immersed in Mexico City. We selected traits that exhibited management potential to provide feasible recommendations for controlling the species. We observed that increasing values of shrub cover, tree cover, and distance to developed areas discouraged the sparrow from invading new sites of the reserve. Simultaneously, greater distances to developed areas promoted the extinction of the species across invaded sites. These effects might be related to resource availability, foraging preferences of the species, predatory exposure, and competition. Preserving tree and shrub cover as a natural barrier for dissuading species arrival represents a key management objective for its control in the reserve. Greater management efforts must be focused at those sites of the reserve that are closer to developed areas, given that the latter may function as source habitats for the House Sparrow. Our approach for identifying management actions that impact the population dynamics of an invasive species might provide crucial results to control this type of populations. Such a strategy could be replicated for other species and natural protected areas to enhance the conservation value of reserves and provide alternatives when dealing with invasive species.


(A) Lack of association between the matrotrophy index per population (MI) and average index of fullness (IFA) for 45 populations of five fish species from the genus Poeciliopsis. (B) Lack of association between the MI per population and the variance of the index of fullness among months (VarIFmonths) for 28 populations of three fish species from the genus Poeciliopsis.
Lack of association between the matrotrophy index per individual (MIind) and the individual index of fullness (IF) for 51 females of five fish species from the genus Poeciliopsis.
Lack of association between the MI per population and average body condition (BCA) for 45 populations of five fish species from the genus Poeciliopsis. Each species is shown in a different panel because of substantial differences among species in the values of body condition (BC). Each symbol represents a different species as per (Figures 1, 2).
Lack of association between the MI per population and the variance of body condition among months (VarBCmonths) for 28 populations of three fish species from the genus Poeciliopsis. Each species is shown in a different panel because of substantial differences among species in the values of BC. Each symbol represents a different species as per (Figures 1, 2).
Negative association between the MIind and individual BC for 51 females of five fish species from the genus Poeciliopsis. Each species is shown in a different panel because of substantial differences among species in the values of BC. Fitted lines represent model-averaged regression coefficients. Each symbol represents a different species as per (Figures 1, 2).
Examination of the Trexler-DeAngelis Model of Maternal Provisioning Reveals That Matrotrophy Is Costly

September 2021

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79 Reads

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4 Citations

The evolution of matrotrophy (post-fertilization maternal provisioning to developing embryos) has been explained through several hypotheses. Trexler and DeAngelis proposed in 2003 a theoretical model that defines the ecological conditions under which matrotrophy would be favored over lecithotrophy (pre-fertilization maternal provisioning). According to this model, matrotrophy offers a selective advantage in environments with abundant and constantly available food, whereas environments with limited and fluctuating food resources should instead promote a lecithotrophic mode of maternal provisioning. This model also proposes that matrotrophy entails the consequence of leaner reproductive females and in turn shorter lifespans. In this study, we examined the Trexler-DeAngelis model using data from 45 populations of five viviparous species from the fish genus Poeciliopsis (family Poeciliidae). We used the matrotrophy index (MI) as a measure of post-fertilization maternal provisioning, and the index of stomach fullness and individual body condition (BC) as proxies for food availability. We also estimated the magnitude of fluctuations in food availability by calculating the temporal variances of these two proxies. Neither abundant nor constantly available food were associated with greater degrees of matrotrophy, which fails to support the predictions of the Trexler-DeAngelis model with respect to the ecological drivers of increased post-fertilization provisioning to embryos. Nonetheless, in all five species we observed that females with greater degrees of matrotrophy had poorer BC compared to females that provided less nutrients to embryos after fertilization. This finding is consistent with one of the expected consequences of advanced matrotrophy according to the Trexler-DeAngelis model, namely, a detriment to the nutritional status of females. Our study provides compelling evidence that gestating females experience a trade-off between post-fertilization provisioning to embryos and self-maintenance, revealing in turn that matrotrophy is a costly reproductive strategy.


Map of the Reserva Ecológica del Pedregal de San Ángel and Universidad Nacional Autónoma de México. A total of 100 observation sites are shown distributed over core, buffer of urban areas. The black area inside Mexico City represents the original extension of the xerophytic scrubland
Effect of environmental covariates on occupancy and detection probabilities. a Effect of shrub cover on the occupancy probability (ψ) during January 2016. b Effect of shrub cover on the occupancy probability (ψ) during May 2016. c Effect of herb cover on the detection probability (p) during May 2016. Error bars and gray lines represent 95% confidence intervals
Effect of shrub cover on the detection probability (p) during May 2016. Error bars and gray lines represent 95% confidence intervals
Analysis of the effect of recreational dog walking on the occupancy probability of the ringtail Bassariscus astutus (Carnivora: Procyonidae) within an urban ecosystem

February 2020

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323 Reads

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5 Citations

Urban Ecosystems

Despite being highly modified environments, cities are important reservoirs of biodiversity. The ecological reserve situated within southern Mexico City houses several species of mammals, including the ringtail Bassariscus astutus. However, the urban areas surrounding the reserve are subject to various types of uses that could affect the presence of wild species inside and out of the reserve areas. Here, I tested the hypothesis that recreational activities, especially the walking of domestic dogs, can cause the ringtails, smaller sized carnivores, to avoid certain areas. To achieve this, I carried out a survey throughout the reserve and urban landscape, and registered the presence of domestic dogs and ringtails by direct observation or via the detection of feces and latrines during two sampling seasons. I used single species occupancy models to determine the environmental covariates associated with the presence of these carnivores within the reserve and the urban areas around it. In addition, I used two-species occupancy models to estimate the species interaction factor between these species to assess if the presence of domestic dogs has a negative effect over the presence of ringtails. My results reveal that shrub cover in the area affects the occurrence of these species, and that there is no clear evidence that domestic dogs negatively affect the occupancy of ringtails during the studied seasons. These results highlight the capacity of the ringtails to adapt to rapid urbanization, but do not guarantee that recreational activities, especially those involving dogs, have no negative effects on the local distribution of other smaller mammals within this urban reserve.


Citations (11)


... Se observó haciendo uso de los árboles, arbustos y matorrales de ambientes urbanos donde hace uso preferencial de las semillas de plantas herbáceas (granívoro) (Zúñiga-Vega et al., 2024). ...

Reference:

Bird diversity in Andean urban ecosystems of the Eastern mountain range of Colombia. A guide for the social appropriation of knowledge / Diversidad de aves urbanas en los Andes Orientales de Colombia: Una guía para la apropiación social de conocimiento
Dynamic occupancy analyses of native birds in an urban ecological reserve reveal seasonal changes in site occupancy and preference for adjacent urbanized areas

Urban Ecosystems

... Thirteen hummingbird species inhabit Mexico City, one of the largest cities in the world (Puga-Caballero et al. 2020, Z uñiga-Vega et al. 2023. This study aims to evaluate how biological factors influence the local distribution of two hummingbird species in a remnant patch of native xerophytic scrubland embedded within the southern part of the city (Zambrano et al. 2016). ...

Raptors in the city: Site occupancy and abundance of a top predator inhabiting urban green spaces within a megacity
  • Citing Article
  • June 2023

Landscape and Urban Planning

... Therefore, it is necessary to test, at the microevolutionary scale, whether the evolution of higher degrees of placentotrophy across populations of a given species promotes greater sexual conflict, which in turn promotes the evolution of enhanced strategies to ensure paternity, similar to the pattern that has been detected at the interspecific scale. The genus Poeciliopsis is characterized by variations between and within species in the degrees of female placentotrophy, where a positive relationship between the amount of maternal supply to developing embryos after fertilization and the degree of placental complexity has been confirmed (Turner 1940;Thibault and Schultz 1978;Reznick et al. 2002;Kwan et al. 2015;Olivera-Tlahuel et al. 2019;Molina-Moctezuma et al. 2020;Saleh-Subaie et al. 2021). Therefore, it is reasonable to hypothesize that, within Poeciliopsis species, males have also experienced phenotypic changes associated with greater post-copulatory sexual selection and sperm competition in populations in which females have evolved higher levels of placentotrophy. ...

Examination of the Trexler-DeAngelis Model of Maternal Provisioning Reveals That Matrotrophy Is Costly

... Based on the above, we modelled the probability of raft use for the first primary period (ψ 1 ), the probability of raft colonization (γ; i.e. the probability of an unused raft at time t becoming used at time t + 1) and raft abandonment (ε), and mink detection probability (p). In order to reduce the effect of uninformative parameters 44 , we developed a limited number of candidate models following a hierarchical four-stage approach 45 . First, we modelled p through a set of models that included each of the observation-level covariates (Table 1), as well as a null model in which no covariates were included (Supplementary Table S1). ...

Identifying management guidelines to control the invasive House Sparrow (Passer domesticus) within natural protected areas through the estimation of local colonization and extinction probabilities

Biological Invasions

... reviews [22,24,27,[32][33][34][35]), they have been poorly studied in urban environments (but see [36][37][38][39][40][41][42][43]). Our limited knowledge of urban migratory bird assemblages indicates that they present lower species richness relative to wildlands [36]. ...

Demographic and spatial responses of resident bird populations to the arrival of migratory birds within an urban environment
  • Citing Article
  • December 2019

Population Ecology

... Es aquí donde encontramos una diferencia al contrastarlo con las respuestas de la encuesta, ya que el cacomixtle norteño fue el mamífero más común con 33 menciones (19 % de las respuestas). Una posible explicación de esto es que el cacomixtle es una especie de hábitos nocturnos para la caza y colecta de sus alimentos(13), lo que le permite evadir a los seres humanos o perros que puedan agredirlo(14). Por consiguiente, obtener una foto convencional de estos ejemplares para reportarlos en el sitio web tiene un mayor grado de dificultad.El 92,5 % de los encuestados afirmaron vivir cerca de áreas verdes; este se puede considerar un factor que genera proximidad entre las especies de fauna urbana y los seres humanos. ...

Analysis of the effect of recreational dog walking on the occupancy probability of the ringtail Bassariscus astutus (Carnivora: Procyonidae) within an urban ecosystem

Urban Ecosystems

... Natural streams have also been identified as key supports for the presence of numerous migratory passerine bird species [12]. Consequently, these spaces acquire special significance as strategic refuges for migratory birds facing the challenges of urban environments for their survival [13]. However, studies on avifauna in riparian areas associated with ephemeral streams in urban settings are scarce [8,9]. ...

Habitat traits that increase the probability of occupancy of migratory birds in an urban ecological reserve
  • Citing Article
  • November 2019

Acta Oecologica

... The reserve comprises 237 ha of conservation area and 48 ha of unprotected scrubland, interspersed with and surrounded by 445 ha of urban landscape that includes: streets, roads, parking lots, buildings, sidewalks and green areas (Fig. 1). This creates a gradient of protection from core reserve and buffer areas to urbanized habitats integrated with the remnant scrubland in which the distribution of the studied species is not restricted to the conservation lands (Zambrano et al. 2019). It is situated between coordinates 19°18 0 31 00 -19°19 0 17 00 N and 99°10 0 20 00 -99°11 0 52 00 W, with an altitude of 2000-2365 m asl (Zambrano et al. 2016, Estañol-Tecuatl & Cano-Santana 2017. ...

Evaluating Socio-Ecological Interactions for the Management of Protected Urban Green Spaces

... It should also be considered that habitats of higher quality are known to be able to support larger numbers of species (Richmond et al. 2010), which may account for the positive interaction observed here, at least in part. In a study of the aggregation of two songbirds (one exotic and the other native), Ramírez-Cruz et al. (2019) concluded that the relationship may be a response to the scarcity of resources, with co-occurrence resulting from aggregations at specific sites with a relative abundance of feeding resources. Varying SIF values have been recorded in other studies of birds, indicating either the independent occurrence of the species (Estevo et al. 2017;Silveira 2018) or species that co-occur less frequently than expected, ie they avoid each other (Haynes et al. 2014;Costa-Neto 2018). ...

This town ain't big enough for both of us…or is it? Spatial co-occurrence between exotic and native species in an urban reserve

... A total of 100 observation sites were placed throughout the REPSA and the surrounding UNAM campus (Fig. 1). Each observation site consisted of a 20-m-radius circle within which we conducted our surveys (Ram ırez-Cruz et al. 2018, 2020. The minimum distance between observation sites was 150 m (Z uñiga- ). ...

How Widespread are Nonnative Species? Estimating Habitat Occupancy in an Ecological Reserve within a Megacity
  • Citing Article
  • January 2018

Natural Areas Journal