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Article

Influence of Natural and Artificial Habitats and Microhabitats on Urban Amphibian Diversity and Behavior

by
Rocio Pamela Demartín
1,*,
Romina Ghirardi
1,2 and
Javier Alejandro López
1,3
1
Grupo de Estudio sobre Anfibios, Laboratorio de Biodiversidad y Conservación de Tetrápodos, Instituto Nacional de Limnología (INALI: CONICET-UNL), Santa Fe 3000, Provincia de Santa Fe, Argentina
2
Instituto de Investigaciones en Ciencias de la Salud, Facultad de Ciencias de la Salud, Universidad Católica de Santa Fe, Santa Fe 3000, Provincia de Santa Fe, Argentina
3
Departamento de Ciencias Naturales, Facultad de Humanidades y Ciencias, Universidad Nacional del Litoral, Santa Fe 3000, Provincia de Santa Fe, Argentina
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(4), 292; https://doi.org/10.3390/d17040292
Submission received: 20 March 2025 / Revised: 16 April 2025 / Accepted: 17 April 2025 / Published: 19 April 2025
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Abstract

:
Species presence in urban landscapes is driven by complex biological and environmental interactions. In this study, we evaluated habitat and microhabitat selection by amphibians in urban environments using correspondence analysis, multiple correspondence analysis, and preference analysis. Data on habitats, microhabitats, and activities were recorded for 26 amphibian species in urban areas. All species were observed in natural habitats, while only 11 in artificial habitats. Leptodactylus latinasus, Leptodactylus macrosternum, Rhinella arenarum, and Rhinella dorbignyi were found in both habitat types, in both aquatic and terrestrial environments. Most individuals (74%) were recorded in natural habitats, predominantly aquatic ones. In artificial habitats (26%), R. arenarum was the most abundant species, primarily using terrestrial habitats. All species exhibited some degree of habitat preference, even generalist species. Amphibian activities were also linked to habitat type, with natural aquatic habitats primarily used for breeding and natural terrestrial habitats for refuge, foraging, and other activities. Our results highlight that heterogeneous natural habitats promote greater species diversity, while artificial habitats restrict amphibian presence. However, the capacity of certain species to adjust to artificial environments underscores the need to enhance these habitats by adding bodies of water, bare ground, and vegetation of all kinds to support the conservation of urban amphibians.

1. Introduction

The rapid growth of cities has profoundly transformed natural landscapes, fragmenting habitats and altering the availability of essential resources for many species [1,2]. This urban expansion not only reduces the extent of natural habitats but also creates environments where built structures and remaining green areas interact in complex ways [3,4,5,6] while introducing a variety of environmental stressors that can substantially alter amphibian assemblages. Habitat fragmentation and spatial isolation constrain dispersal and gene flow [7,8], increased competition or predation modifies interspecific interactions, and elevated exposure to pollutants and ecological traps impacts individual health and survival [9,10,11]. Moreover, habitats and microhabitats are altered by shifts in microclimate, elevated levels of noise and light pollution, and the degradation or loss of critical sites for breeding, shelter, and foraging [12,13,14,15]. Collectively, these factors drive changes in amphibian community structure, acting as environmental filters [16]. The presence of a species within this landscape is shaped by multiple biological and environmental factors, and its survival depends on its ability to adapt to changing conditions and exploit available resources [17,18,19,20].
Amphibians are ectothermic organisms with permeable skin that rely directly on environmental conditions to regulate their body temperature and moisture levels [21,22]. Their survival depends on access to humid microhabitats that prevent dehydration and buffer temperature fluctuations [23]. However, urbanization modifies these environmental conditions, often increasing temperatures and reducing the availability of humid refuges. Additionally, it alters habitat structure and quality, leading to habitat loss, fragmentation, and degradation [24,25,26,27,28]. Urban environments contain a mosaic of habitats that can be broadly classified as natural or artificial, depending on their origin, and these in turn, depending on their hydrological characteristics, as aquatic or terrestrial. While aquatic habitats are essential for breeding and larval development, terrestrial habitats provide shelter, foraging opportunities, and connectivity between water bodies and other suitable areas [29]. Within each habitat, amphibians select specific microhabitats that offer the environmental conditions necessary for different activities. Furthermore, in urban environments, heat retained by structures such as buildings, sidewalks, and streets creates warmer microclimates than those found in natural areas, altering the characteristics of available microhabitats [30,31]. This phenomenon, known as the urban heat island effect, can increase surface temperatures and reduce relative humidity, limiting the availability of cool, humid microhabitats that amphibians need to prevent dehydration and thermal stress [32,33]. The reduction in humid areas, such as water bodies, saturated soils, or shaded zones, is a key constraint, as amphibians require high moisture levels to prevent dehydration, facilitate gas exchange through their skin, and support oviposition and larval development [29]. Additionally, amphibians select microhabitats that provide shelter from predators and offer optimal conditions to minimize competition and exposure to pathogens [34,35]. In this context, the texture and composition of urban surfaces, such as pavement, artificial grass, or compacted soils, influence their capacity to retain water and provide adequate refuge [36]. The accessibility of these microhabitats also depends on landscape permeability, meaning the ability to move through the environment without encountering natural or artificial barriers. All these factors interact to shape species’ presence in a given area and determine their ability to adapt to changes in the urban landscape [37].
In order to provide insights into amphibian responses to landscape changes and also offer key information for developing management and conservation strategies that mitigate the negative impacts of urban expansion and promote biodiversity coexistence within cities [38], our objective was to analyze the use and preference of habitats and microhabitats by amphibians to understand how they interact with the surrounding landscape and persist in urban environments.

2. Materials and Methods

2.1. Study Area

The survey was carried out in the metropolitan area of Santa Fe City, including Santa Fe and San José del Rincón cities, and Arroyo Leyes commune (Department La Capital, Province of Santa Fe, Argentina) (Figure 1). Santa Fe City (31°42′ S; 60°46′ W) is the administrative and political capital of the Province of Santa Fe. It is the second-largest urban center of the province and has around 425,000 inhabitants according to data available from the last census [39]. Santa Fe City is situated in the Middle Paraná River floodplain, surrounded by the Salado River to the west and south and the Paraná River fluvial system to the east and south. San José del Rincón City (31°36′ S; 60°34′ W) has a population of more than 11,000 inhabitants [39] and borders Arroyo Leyes commune to the north and Santa Fe City to the south. It is located 13 km north of the urban center of Santa Fe City. Arroyo Leyes commune (31°35′ S; 60°33′ W) has a population of almost 3300 inhabitants [39], borders San José del Rincón City to the south, and is located 18 km north of the urban center of Santa Fe City. The three localities are situated within the Espinal ecoregion, characterized by a mix of forests and savannas, as well as the Paraná Delta and Islands ecoregion, encompassing the floodplain along the Middle Paraná River [40]. These areas are positioned at altitudes below 25 m above sea level, exhibiting a thermal amplitude between 31 °C and 34 °C [41] and an average annual rainfall of 1020.60 mm registered during the last 10 years (2015–2024) by the Centro de Informaciones Meteorológicas of the Facultad de Ingeniería y Ciencias Hídricas of the Universidad Nacional del Litoral (CIM: FICH-UNL).

2.2. Sampling Methods

Field surveys were conducted during the spring and summer months (October–March) of 2021, 2022, 2023, and 2024. A total of 70 surveys were carried out across 14 selected urban green sites, with five sampling events per site. Each survey included active searches along two transects (50 m × 2 m) per site, lasting 20 min each. Amphibians were identified to species level at the time of observation. For each individual, we recorded the habitat, microhabitat (substrate), and activity performed at the time of observation. Habitats were classified into two main categories: natural habitats (including green areas, ditches, vegetation patches, and bodies of water with natural sediment bottoms) and artificial habitats (streets, sidewalks, sports courts, buildings, canals, and artificial pools with impermeable bottoms). Each category was further divided into aquatic and terrestrial habitats, where aquatic habitats were defined as areas covered by at least 1 cm-deep water film. Considering substrates where amphibians were registered, we identified ten microhabitat types: open water, grasses, leaf litter, ant nests, woody substrates, floating macrophytes, emergent macrophytes, bare ground (sand/soil), bromeliads, and impermeable surfaces (cement, plastic waste, metals, bricks). Observed activities were categorized into: (1) foraging (individuals feeding or searching for prey), (2) sheltering (hidden individuals), (3) reproduction (calling, amplexus, or mate selection), and (4) other activities (movement without clear purpose, resting, thermoregulating, hydrating, etc.).
Analyses were conducted to assess associations between categorical variables: species, habitat, microhabitat, and activity. To obtain a visual representation of these associations, a general correspondence analysis (CA) was performed to examine the relationship between species and habitat types. Additionally, two multiple correspondence analyses (MCAs) were conducted separately for natural and artificial habitats, incorporating all four studied variables. Variables were color-coded based on their cos² values. The cos² is obtained from the decomposition of the total inertia and represents the proportion of a category’s variance explained by a given dimension that indicates the quality of representation of each variable within the dimensions of the plot [42]. To calculate species distribution preference and assess whether it varied according to activity, we used the weighted preference index (WPI) proposed by Clark et al. (1999) [43]. This index is based on the standard deviation of proportions across samples, weighted by the number of individuals observed in each category, and is calculated as follows:
W P I j = ∑ i = 1 s p i ( n i j p i − N j P ) 2 P
where i represents habitat type up to S (total number of habitat types), pi is the number of individuals present in habitat i, P is the total number of individuals observed, nij is the number of individuals of species j in habitat i, and Nj is the total number of individuals of species j.
The WPI takes a value of 0 when a species exhibits completely generalist behavior (i.e., it is evenly distributed across all habitat types) and increases as its distribution deviates from random [43].
To assess the significance of the observed values, we performed a resampling analysis based on the number of individuals per species and compared the results against a null distribution, using the 95th percentile as the significance threshold. This analysis was conducted at two levels of resolution: first at the habitat level (n = 4), and then at the microhabitat level (n = 23), considering all possible observed combinations of habitat and microhabitat.
Additionally, to explore whether habitat preference was associated with species activities, we repeated the analysis considering each species individually. In this case, Nj represents the number of individuals of species j, and nij corresponds to the number of individuals of species j performing activity i in a given site.
All analyses were conducted using R software [44] employing the following packages: FactoMineR [45], factoextra [46], tidyverse [47], lme4 [48], and mgcv [49].

3. Results

A total of 26 amphibian species belonging to 12 genera and 6 families were recorded (Table 1), all listed with the threat category of least concern by the International Union for Conservation of Nature [50]. All species were present in natural habitats, whereas only 11 species were found in artificial habitats (Table 2). The species Leptodactylus latinasus, Leptodactylus macrosternum, Rhinella arenarum, and Rhinella dorbignyi utilized both natural and artificial habitats, in both aquatic and terrestrial environments. All other species were restricted to fewer combinations, occurring only in natural or artificial terrestrial habitats, or in just one type of environment.
All microhabitat categories were present in natural habitats, whereas only five categories were observed in artificial habitats: open water, grasses, leaf litter, bare ground, and impermeable surfaces. The species recorded in the greatest number of microhabitats (n = 8) were Dendropsophus nanus (exclusively in natural habitats), and Boana raniceps and Rhinella arenarum (both in natural and artificial habitats).
In contrast, Physalaemus biligonigerus, Physalaemus santafecinus, Pseudopaludicola falcipes, Boana punctata, Pseudis platensis, Scinax squalirostris, and Trachycephalus thyponius were found in only one microhabitat, all within natural habitats.
The majority of observed individuals (74%) were found in natural habitats, with a higher presence in aquatic environments (61%). In natural aquatic habitats, 24 species were recorded, with the open water microhabitat being the most utilized (38% of individuals and 17 species). In natural terrestrial habitats, the most frequently used microhabitat was grasses (51% of individuals and 15 species) (Table 3). On the other hand, 26% of individuals were found in artificial habitats, with 64% occupying terrestrial environments. The species Rhinella arenarum accounted for 89% of the total abundance, while the remaining 11% was distributed among 10 species from various genera. In artificial aquatic habitats, four species were recorded, whereas eleven species were observed in artificial terrestrial habitats, with more than 80% of individuals found on impermeable surfaces (Table 4).
The correspondence analysis (Figure 2) revealed associations between species and habitat types, explaining 99.3% of the total variance in the first two dimensions. In sum, 10 species were strongly associated with natural aquatic habitats, while 14 species were moderately associated with natural terrestrial habitats. In contrast, R. arenarum was primarily associated with artificial habitats. The multiple correspondence analysis of the natural habitat (Figure 3) explained 12% of the total variance in the first two dimensions. In the natural aquatic habitat, the species D. nanus was associated with the floating macrophyte microhabitat, while R. dorbignyi was linked to the water microhabitat, both related to reproductive activity. In the natural terrestrial habitat, the grass microhabitat was associated with other activities. The multiple correspondence analysis of the artificial habitat (Figure 4) explained 18.9% of the total variance in the first two dimensions, so we used the best-represented variables to interpret the graph. The species L. limellum and L. mystacinus were associated with the bare ground microhabitat and sheltering activity. On the other hand, R. arenarum was linked to impermeable surfaces, while L. macrosternum was associated with the water microhabitat, both related to reproductive activity and artificial aquatic habitats.
The preference analyses allowed us to assess whether species select specific habitats and microhabitats, both in general and in relation to the different recorded activities. Only species with more than 10 observations could be included in the analysis; therefore, habitat preference results are presented for 15 species, as the remaining 11 species had low observation counts.
Regardless of activity (Figure 5), all species exhibited at least minimal habitat and microhabitat selection. Regarding habitat selection, L. macrosternum and Pithecopus azureus showed the lowest selectivity, whereas D. nanus and R. arenarum demonstrated higher selectivity than the other species. In terms of microhabitat selection, Leptodactylus gracilis and B. raniceps exhibited the lowest selectivity, while D. nanus and R. arenarum again displayed the highest microhabitat selectivity (Table A1).
Considering activity, significant preference indices indicated that natural aquatic habitats were selected for reproductive activities, whereas natural terrestrial habitats were preferred for foraging, sheltering, and other activities (Figure 6). Only two species selected artificial habitats: R. arenarum for sheltering and reproductive activities, and Scinax nasicus for other activities. Preferred microhabitats included water for reproduction, bare ground for sheltering, grass for other activities, and bare ground and anthills for foraging. Preference analyses indicated that Rhinella arenarum selected both natural and artificial habitats, as well as multiple microhabitats for reproductive and sheltering activities.

4. Discussion

The results highlight that natural habitats, along with the diversity of microhabitats they contain, support a greater species diversity compared to artificial habitats. This may be attributed to the fact that natural habitats tend to offer a variety of resources to meet the needs of feeding, reproduction, and shelter [51]. In contrast, artificial habitats limit the diversity of species that can persist in them, as they fail to provide the appropriate substrate to meet basic needs [52]. Although all recorded species showed some degree of tolerance to urban environments, differences were observed in how they use available habitats and microhabitats. These differences may be associated with varying levels of ecological plasticity, both across life stages and at the habitat or microhabitat scale in adults [15]. This reinforces the importance of considering species-specific responses to different urbanization scenarios. Some species exhibit a remarkable capacity to use both natural and artificial habitats in urban environments, demonstrating high ecological plasticity. The species Leptodactylus latinasus, Leptodactylus macrosternum, Rhinella arenarum, and Rhinella dorbignyi stood out for their presence in all four evaluated habitat types, highlighting their ecological plasticity. Their ability to exploit both natural and artificial environments suggests that they are habitat generalists. At the same time, their presence in urban areas in cities of different regions reflects their tolerance to urbanization [53,54,55]. These characteristics allow them to survive in anthropogenic landscapes and carry out various activities, taking advantage of different resources available in cities.
The species R. arenarum and Scinax nasicus, which have been classified as urban-tolerant by various studies [54,55,56,57], were the only ones that preferred artificial habitats for specific activities such as sheltering and reproduction. Their preference for urban sites could be explained by lower interspecific competition [58], the abundance of food associated with urban lighting ([59], personal observation), the presence of ephemeral water bodies with few predators [60] where they invest significant energy in reproduction (personal observation), and the availability of shelter in cracks within buildings, lighting fixtures, and artificial structures, which seem essential for survival and success in fragmented environments ([61], personal observation).
The three species associated with the highest number of microhabitats exhibited notable ecological plasticity, demonstrating a high capacity to establish themselves in different environments. However, it was observed that all microhabitats occupied by Dendropsophus nanus were natural habitats, whereas Boana raniceps and R. arenarum utilized microhabitats in both natural and artificial habitats. The restriction of D. nanus to natural habitats could be related to its small body [62], which limits its dispersal capacity and makes it more susceptible to desiccation in more exposed environments [63]. Therefore, D. nanus can be considered a microhabitat generalist, but not a habitat generalist, whereas B. raniceps and R. arenarum can be classified as generalists in both habitat and microhabitat use.
However, some species such as Elachistocleis bicolor, R. arenarum, R. dorbignyi, D. nanus and S. nasicus that exhibited generalist behavior in habitat or microhabitat use also showed preference patterns for specific habitats or microhabitats, as reflected in their selectivity indices compared to other species. This apparent contrast suggests that although these species can occupy diverse environments, they may still prefer particular habitats or microhabitats that provide optimal conditions for survival [20,64].
On the other hand, species observed in only one microhabitat may be influenced by different factors, such as habitat specificity, shifts in microhabitat use to survive or exploit resources in urban environments, or biases in recorded activities. For instance, Boana punctata was recorded exclusively in floating macrophytes while performing different activities, which could indicate a true preference for this microhabitat [65]. Similarly, Physalaemus santafecinus was always observed in the water microhabitat during reproductive activity. While P. santafecinus typically exhibits both aquatic and terrestrial habits [62], its lifestyle in urban sites may shift toward greater dependence on water, not only for reproduction but also for other activities that are difficult to observe in the water, potentially helping to avoid physiological stress associated with urban environments [66,67]. However, this hypothesis should be corroborated in the field. Otherwise, it is possible that the use of other microhabitats for different activities was not recorded. The remaining species, observed in only one type of microhabitat, were represented by a low number of individuals, which limits our possibility to draw strong inferences regarding their microhabitat preferences. However, based on the literature, we found that our field observations are consistent with previously reported microhabitat use for several of these species. For instance, Leptodactylus mystacinus and Pseudopaludicola falcipes has been recorded on bare soil [68]; Leptodactylus podicipinus in leaf litter, bare soil, and open water [69]; Pseudis platensis in open water [69]; Scinax squalirostris among emergent macrophytes [68,69]; and Trachycephalus typhonius on trees [69,70]. It is important to note that none of these studies was conducted in urban environments; therefore, these species are consistent with their selection of microhabitats in natural and urban environments.
Regarding habitat and microhabitat use for different activities, as expected, most species selected water bodies for reproductive activity. This microhabitat is essential for completing key amphibian life cycle processes, such as egg-laying and larval development [71,72]. Even shallow water bodies (>1 cm depth) have proven to be crucial for the persistence of populations in modified urban landscapes [73].
The preference for bare ground as a refuge microhabitat among terrestrial species is likely due to its thermal stability, higher moisture retention, and lower exposure to predators [74,75]. Similarly, grasses, which were selected for various other activities, may provide a dense vegetation cover that allows amphibians to move with reduced predator exposure, prevent desiccation, and even forage on invertebrates present in this microhabitat [76,77].
Foraging activity, which was significant in different microhabitats across various species, may be explained by species-specific dietary preferences. For instance, E. bicolor, an ant specialist with an active foraging strategy [78,79], showed a preference for ant mound microhabitats, reinforcing the relationship between its feeding strategy and the selection of specific resources. Additionally, Physalaemus albonotatus and R. dorbignyi exhibited preferences for ant mounds and bare ground, respectively. Both species have generalist and intermediate diets and display opportunistic feeding behaviors [80,81,82]. Their diet consists largely of ants [82,83,84], aligning with their habitat preferences and the ease of prey detection on bare ground (personal observation of R. dorbignyi feeding on an ant trail). This suggests that foraging site selection is influenced by specific habitat characteristics and temporal factors, allowing adaptation to prey availability in the environment [85].
Notably, Rhinella arenarum showed a preference for multiple habitats and microhabitats for reproductive and refuge activities, highlighting its remarkable ecological plasticity and ability to thrive in highly fragmented environments [54]. A particularly relevant aspect is its inclination toward artificial sites with impermeable surfaces, where populations exceeding 100 individuals have been observed, most engaging in reproductive activities. This preference could be explained by the favorable conditions of these ephemeral pools, where reduced competition and predator presence [59,61] combined with increased reproductive success opportunities [86] minimize risks and facilitate larval development. From a phylogenetic perspective, such ecological flexibility is consistent with patterns observed in other bufonids, both in South America and globally, which have been documented in urban habitats, exhibiting a notable capacity to persist in human-modified environments [41,87,88,89,90,91,92]. This capacity may be linked to general traits shared by the family, such as wide geographic distribution, resistant skin, and generalist feeding strategies, among others, which collectively may promote their success in urban environments [67,93,94,95,96].
Nevertheless, such plasticity does not exempt bufonids—or other amphibian species—from the multiple pressures encountered in urban environments. A variety of factors may influence the survival and behavior of urban amphibians. Urban predators, including insects, fish, reptiles, birds, and domestic mammals observed at the sampled sites, likely exert constant pressure throughout all stages of the amphibian life cycle [97,98]. Additionally, roadkill-related mortality in areas fragmented by road infrastructure suggests a direct effect of urbanization on individual survival [99]. Ecological traps—such as artificial water bodies with smooth edges—may also pose particular risks, as they often fail to accommodate the ecological requirements of local fauna [9]. Within this context, we observed the predation of an adult R. arenarum by an aquatic turtle, illustrating the potential hazards present in such environments. Although environmental factors such as artificial lighting and acoustic pollution were recorded, no clear effects on individual activity were detected. However, previous studies have reported that such conditions can affect calling behavior, activity patterns, and acoustic communication in anurans [100,101,102], and their potential influence should not be overlooked. Finally, anthropogenic impacts such as waste accumulation, vegetation removal, pesticide application, and wastewater discharge were observed—some coinciding with the presence of injured or dead individuals—which aligns with previously reported findings [103,104]. While no direct causal relationships could be established, these observations underscore the need for more targeted research on the impacts of local disturbances on the health of urban amphibians [105]. Understanding the ecological tolerance limits of amphibians, as well as the mechanisms that enable their persistence in urban landscapes, is therefore essential for developing effective conservation strategies in these environments.

5. Conclusions

Our results highlight the importance of natural habitats for the conservation of amphibian diversity in urban environments, as they host a greater variety of species and microhabitats compared to artificial habitats. However, some species, such as Rhinella arenarum and Scinax nasicus, have demonstrated remarkable ecological plasticity, adapting to urban environments by selecting specific microhabitats for key activities such as shelter and reproduction. The ability of certain species to persist in modified landscapes suggests that the availability of suitable structural elements, such as temporary water bodies and refuges in artificial surfaces, may favor their survival. Nevertheless, habitat fragmentation and the reduction in areas with optimal moisture conditions and vegetation cover continue to pose challenges for most species. In this context, the design of urban planning strategies that integrate green spaces with suitable microhabitats could mitigate the effects of urbanization on amphibian communities and promote their coexistence in urban environments.

Author Contributions

Conceptualization, R.P.D., R.G., and J.A.L.; methodology, R.P.D., R.G., and J.A.L.; software, R.P.D.; validation, R.P.D., R.G., and J.A.L.; formal analysis, R.P.D.; investigation, R.P.D., R.G., and J.A.L.; resources, R.G., and J.A.L.; data curation, R.P.D.; writing—original draft preparation, R.P.D.; writing—review and editing, R.P.D., R.G., and J.A.L.; visualization, R.P.D., R.G., and J.A.L.; supervision, R.G., and J.A.L.; project administration, R.G., and J.A.L.; funding acquisition, R.G., and J.A.L. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Fondo para la Investigación Científica y Tecnológica of Agencia Nacional de Promoción de la Investigación, el Desarrollo Tecnológico y la Innovación (PICT2020-Serie A-03835).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author(s).

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

Abbreviations

The following abbreviations are used in this manuscript:
DOAJDirectory of Open Access Journals
TLAthree-letter acronym
LDlinear dichroism

Appendix A

Table A1. Habitat and microhabitat significantly selected by species. Significance was determined by observing that simulated indices were greater than the 95th quantile of the null distribution and that the observed abundance for each species exceeded the abundance from resampling.
Table A1. Habitat and microhabitat significantly selected by species. Significance was determined by observing that simulated indices were greater than the 95th quantile of the null distribution and that the observed abundance for each species exceeded the abundance from resampling.
SpeciesHabitatMicrohabitat
Scinax nasicusNatural terrestrialNatural aquatic open water
Natural terrestrial bromeliad
Natural terrestrial woody substrates
Rhinella dorbignyiNatural aquaticNatural aquatic open water
Natural aquatic grasses
Natural aquatic bare ground
Rhinella arenarumArtificial aquaticArtificial aquatic impermeable surface
Artificial terrestrialArtificial terrestrial impermeable surface
Pithecopus azureus Natural aquatic emergent macrophyte
Natural terrestrial emergent macrophyte
Physalaemus albonotatusNatural terrestrialNatural terrestrial grasses
Leptodactylus macrosternumNatural terrestrialArtificial aquatic water
Natural aquatic grasses
Natural terrestrial grasses
Natural terrestrial bare ground
Leptodactylus luctatorNatural terrestrialNatural aquatic bare ground
Natural terrestrial grass
Natural terrestrial leaf litter
Natural terrestrial bare ground
Lysapsus limellumArtificial terrestrialArtificial terrestrial bare ground
Natural aquatic open water
Leptodactylus latinasusNatural terrestrialNatural aquatic bare ground
Natural terrestrial grasses
Natural terrestrial impermeable surface
Leptodactylus gracilisNatural terrestrialNatural aquatic open water
Natural terrestrial bare ground
Elachistocleis bicolorNatural terrestrialNatural aquatic open water
Natural terrestrial grasses
Natural terrestrial ant nest
Natural terrestrial bare ground
Dendropsophus sanborniNatural aquaticNatural aquatic bromeliad
Natural aquatic floating macrophyte
Natural aquatic emergent macrophyte
Dendropsophus nanusNatural aquaticNatural aquatic woody vegetation
Natural aquatic floating macrophyte
Natural aquatic emergent macrophyte
Boana ranicepsNatural terrestrialArtificial terrestrial bare ground
Natural terrestrial woody vegetation
Boana pulchellaNatural terrestrialNatural terrestrial woody vegetation

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Diversity 17 00292 g001
Figure 1. Satellite image of the metropolitan area of Santa Fe City (study area). The location of the study area at the scale of South America is shown on the right. Urban areas are represented in gray.
Figure 1. Satellite image of the metropolitan area of Santa Fe City (study area). The location of the study area at the scale of South America is shown on the right. Urban areas are represented in gray.
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Figure 2. Correspondence analysis relating species to the habitat used at the time of observation.
Figure 2. Correspondence analysis relating species to the habitat used at the time of observation.
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Figure 3. Multiple correspondence analysis relating species to the microhabitat used and the activity performed at the time of observation in the natural habitat. The color of the names indicates the cos² value, reflecting the quality of representation of each species or category in the selected dimensions. Higher values (red, yellow, orange) indicate better representation, while lower values (light to dark green) suggest greater contribution in other, non-visualized dimensions.
Figure 3. Multiple correspondence analysis relating species to the microhabitat used and the activity performed at the time of observation in the natural habitat. The color of the names indicates the cos² value, reflecting the quality of representation of each species or category in the selected dimensions. Higher values (red, yellow, orange) indicate better representation, while lower values (light to dark green) suggest greater contribution in other, non-visualized dimensions.
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Figure 4. Multiple correspondence analysis relating species to the microhabitat used and the activity performed at the time of observation in the artificial habitat. The color of the names indicates the cos² value, reflecting the quality of representation of each species or category in the selected dimensions. Higher values (red, yellow, orange) indicate better representation, while lower values (dark green and dark) suggest greater contribution in other, non-visualized dimensions.
Figure 4. Multiple correspondence analysis relating species to the microhabitat used and the activity performed at the time of observation in the artificial habitat. The color of the names indicates the cos² value, reflecting the quality of representation of each species or category in the selected dimensions. Higher values (red, yellow, orange) indicate better representation, while lower values (dark green and dark) suggest greater contribution in other, non-visualized dimensions.
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Figure 5. Species preferences for habitat and microhabitat. Higher z-values indicate greater habitat and microhabitat selectivity.
Figure 5. Species preferences for habitat and microhabitat. Higher z-values indicate greater habitat and microhabitat selectivity.
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Figure 6. Species habitat and microhabitat preferences according to the activity performed at the time of observation. NT: natural terrestrial habitat: NA: natural aquatic habitat; AT: artificial terrestrial habitat; AA: artificial aquatic habitat; flt. macrophyte: floating macrophyte; emg. macrophyte: emergent macrophyte; imp. surface: impermeable surface.
Figure 6. Species habitat and microhabitat preferences according to the activity performed at the time of observation. NT: natural terrestrial habitat: NA: natural aquatic habitat; AT: artificial terrestrial habitat; AA: artificial aquatic habitat; flt. macrophyte: floating macrophyte; emg. macrophyte: emergent macrophyte; imp. surface: impermeable surface.
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Table 1. Family, genus, species, and common name in English (sensus Frost, 2024) and local common name in Spanish (sensus Ghirardi and López 2020) of amphibians of the order Anura recorded in urban sites within the metropolitan area of Santa Fe City, Argentina.
Table 1. Family, genus, species, and common name in English (sensus Frost, 2024) and local common name in Spanish (sensus Ghirardi and López 2020) of amphibians of the order Anura recorded in urban sites within the metropolitan area of Santa Fe City, Argentina.
Families (6)Genera (12)Species (26)Common NameLocal Common Name
BufonidaeRhinellaR. arenarumArgentine toadSapo común
R. dorbingyiD’Orbigny’s toadSapo de panza amarilla
HylidaeBoanaB. pulchellaMontevideo tree frogRana del zarzal
B. punctataPolka-dot tree frogRana punteada
B. ranicepsChaco tree frogRana trepadora chaqueña
DendropsophusD. nanusDwarf tree frogRana enana
D. sanborniSanborn’s tree frogRana enana de Sanborni
PseudisP. limellumUruguay harlequin frogRana nadadora chica
P. platensisParadoxical frogRana paradoxa
ScinaxS. acuminatusMato Grosso snouted tree frogRana hocicuda chaqueña
S. nasicusLesser snouted tree frogRana de los baños
S. squalirostrisStriped snouted tree frogRana trepadora rayada
TrachycephalusT. typhoniusRana Lechera ComunRana lechosa
LeptodactylidaeLeptodactylusL. gracilisDumeril’s striped frogRana rayada
L. latinasusOven frogRana piadora
L. luctatorWrestler frogRana criolla
L. macrosternumMiranda’s white-lipped frogRana chaqueña
L. mystacinusMustached frogRana de bigotes
L. podicipinusPointedbelly frogRana puntiaguda espumera
PhysalaemusP. albonotatusMenwig frogRana llorona
P. biligonigerusWeeping frogRana llorona
P. santafecinusHelvetia dwarf frogRana llorona
PseudopaludicolaP. falcipesHensel’s swamp frogRana enana de hensel
MicrohylidaeElachistocleisE. bicolorTwo-colored oval frogRana aceituna
OdontophrynidaeOdontophrynusO. americanusCommon lesser escuerzoEscuerzo común
PhyllomedusidaePithecopusP. azureusEarless monkey leaf frogRana mono
Table 2. Number (percentage) of individuals of each amphibian species recorded in artificial and natural habitats, categorized by aquatic and terrestrial environments, in the metropolitan area of Santa Fe City, Argentina.
Table 2. Number (percentage) of individuals of each amphibian species recorded in artificial and natural habitats, categorized by aquatic and terrestrial environments, in the metropolitan area of Santa Fe City, Argentina.
SpeciesArtificialNatural
AquaticTerrestrialTotalAquaticTerrestrialTotal
R. arenarum157 (30)245 (47)402 (77)25 (5)98 (18)123 (23)
R. dorbignyi1 (0.6)3 (2)4 (2.6)140 (88)15 (9.4)155 (97.4)
B. pulchella01 (6)1 (6)3 (17)14 (77)17 (94)
B. punctata0009 (100)09 (100)
B. raniceps03 (16)3 (16)5 (26)11 (58)16 (84)
D. nanus000300 (95)15 (5)315 (100)
D. sanborni00070 (93)5 (7)75 (100)
P. limellum08 (35)8 (35)15 (65)015 (65)
P. platensis0001 (100)01 (100)
S. acuminatus00002 (100)2 (100)
S. nasicus03 (3)3 (3)29 (32)58 (65)87 (97)
S. squalirostris0001 (100)01 (100)
T. thyponius0002 (100)02 (100)
L. gracilis00014 (36)24 (64)38 (100)
L. latinasus1 (2.3)3 (7)4 (9.3)8 (18)32 (72.7)40 (90.7)
L. luctator06 (10)6 (10)16 (27)38 (63)54 (90)
L. macrosternum5 (6)10 (12)15 (18)33 (40)34 (42)67 (82)
L. mystacinus03 (37)3 (37)1 (13)4 (50)5 (63)
L. podicipinus0008 (89)1 (11)9 (100)
P. albonotatus00012 (21)45 (79)57 (100)
P. biligonigerus00002 (100)2 (100)
P. santafecinus0005 (100)05 (100)
P. falcipes0001 (100)01 (100)
E. bicolor01 (0.7)1 (0.7)60 (39.3)92 (60)152 (99.3)
O. americanus0001 (33)2 (67)3 (100)
P. azureus00012 (67)6 (33)18 (100)
Table 3. Number (percentages) of individuals of each amphibian species recorded in natural habitats, categorized by microhabitats, in the metropolitan area of Santa Fe City, Argentina. Flt. Macro.: floating macrophyte, Emg. Macro: emergent macrophyte, woody subs.: woody substrate, Imp. Surf.: impermeable surface.
Table 3. Number (percentages) of individuals of each amphibian species recorded in natural habitats, categorized by microhabitats, in the metropolitan area of Santa Fe City, Argentina. Flt. Macro.: floating macrophyte, Emg. Macro: emergent macrophyte, woody subs.: woody substrate, Imp. Surf.: impermeable surface.
SpeciesFlt.
Macro.		Emg. Macro.Bare GroundOpen WaterGrassesLeaf LitterWoody Subs.BromeliadsImp. Surf.Ant Nests
R. arenarum0024 (20)16 (13)78 (63)05 (4)000
R. dorbignyi0010 (6)122 (79)23 (15)00000
B. pulchella2 (12)1 (6)1 (6)00013 (76)000
B. punctata9 (100)000000000
B raniceps2 (13)3(19)4 (25)01 (6)1 (6)5 (31)000
D. nanus243 (77)51 (16)1 (0.3)1 (0.3)11 (3)1 (0.3)6 (2)1 (0.4)00
D. sanborni58 (77)11(15)003 (4)02 (3)1 (1)00
P. limellum1 (7)0014 (93)000000
P. platensis0001 (100)000000
S. acuminatus001 (50)000001 (50)0
S. nasicus006 (7)23 (26)9 (10)047 (54)2 (2)00
S. squalirostris01 (100)00000000
T. thyponius02 (100)00000000
L. gracilis0018 (47)14 (37)6 (16)00000
L. latinasus009 (23)5 (13)25 (62)0001 (2)0
L. luctator3 (6)020 (37)8 (15)21 (39)2 (4)0000
L. macrosternum2 (3)012 (18)13 (19)40 (60)00000
L. mystacinus003 (60)1 (20)1 (20)00000
L. podicipinus003 (33)5 (56)01 (11)0000
P. albonotatus008 (14)8 (14)40 (70)00001 (2)
P. biligonigerus00002 (100)00000
P. santafecinus0005 (100)000000
P. falcipes001 (100)0000000
E. bicolor0036 (24)57 (37)50 (33)2 (1)1 (1)006 (4)
O. americanus0001 (33)2 (67)00000
P. azureus017 (94)01 (6)000000
Table 4. Number (percentage) of individuals of each amphibian species recorded in artificial habitats, categorized by microhabitats, in the metropolitan area of Santa Fe City, Argentina.
Table 4. Number (percentage) of individuals of each amphibian species recorded in artificial habitats, categorized by microhabitats, in the metropolitan area of Santa Fe City, Argentina.
SpeciesGrassesLeaf LitterBare GroundImpermeable SurfaceOpen Water
R. arenarum1 (0.2)2 (0.5)16 (4)383 (95.3)0
R. dorbignyi001 (25)3 (75)0
B. pulchella001 (100)00
B. raniceps002 (67)1 (33)0
P. limellum008 (100)00
S. nasicus0003 (100)0
L. latinasus001 (25)3 (75)0
L. luctator1 (17)01 (17)4 (67)0
L. macrosternum01 (7)011 (73)3 (20)
L. mystacinus003 (100)00
E. bicolor0001 (100)0
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Demartín, R.P.; Ghirardi, R.; López, J.A. Influence of Natural and Artificial Habitats and Microhabitats on Urban Amphibian Diversity and Behavior. Diversity 2025, 17, 292. https://doi.org/10.3390/d17040292

AMA Style

Demartín RP, Ghirardi R, López JA. Influence of Natural and Artificial Habitats and Microhabitats on Urban Amphibian Diversity and Behavior. Diversity. 2025; 17(4):292. https://doi.org/10.3390/d17040292

Chicago/Turabian Style

Demartín, Rocio Pamela, Romina Ghirardi, and Javier Alejandro López. 2025. "Influence of Natural and Artificial Habitats and Microhabitats on Urban Amphibian Diversity and Behavior" Diversity 17, no. 4: 292. https://doi.org/10.3390/d17040292

APA Style

Demartín, R. P., Ghirardi, R., & López, J. A. (2025). Influence of Natural and Artificial Habitats and Microhabitats on Urban Amphibian Diversity and Behavior. Diversity, 17(4), 292. https://doi.org/10.3390/d17040292

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