Cities worldwide are expanding in area and human population, posing multiple challenges to amphibian populations, including habitat loss from removal of wetlands and terrestrial upland habitat, habitat fragmentation due to roads and the built environment, and habitat degradation from pollutants, extensive human use and introduced species.
We developed an eight-step urban amphibian conservation framework based on established monitoring, analytical methods and community engagement to enable amphibian conservation in a large urban centre. The framework outlines a process used to conserve biodiversity in a complex landuse and decision-making environment supported by a series of successive complementary modelling techniques to measure amphibian presence, priority habitat and functional connectivity.
We applied the framework in Calgary, Alberta, Canada to illustrate its potential. Here, urbanization has reduced wetlands by 90% and ecological knowledge on amphibians was poor. We improved knowledge on amphibian diversity and distribution, identified core wetlands and movement pathways for amphibian species and identified barriers in the wetland network where construction or restoration measures could re-establish amphibians or increase their densities. This knowledge was shared with ecologists and city planners for implementation through appropriate policies and plans.
Our framework provides a series of stepwise products to improve an urban municipality's ability to restore or conserve priority habitat and movement pathways necessary for amphibian survival under pressure from multiple land uses. The framework provides a platform to identify city plans, policy and or programmes and to derive necessary information to support amphibian conservation.

1 INTRODUCTION

Biodiversity is an integral component of ecosystem services and an important indicator of human well-being. Half of the world's population lives in cities, increasing the need to understand and address impacts on urban biodiversity (Zari, 2018). Urbanization is a complex process where people and associated built infrastructure alter natural hydrological, physical and chemical processes, often resulting in changes to animal and plant communities (McKinney, 2006).

Urbanization profoundly impacts ecological ecosystems through an increase in impervious surfaces and pollutants resulting in habitat degradation, loss and fragmentation for much native biodiversity. As cities adopt mandates to protect, maintain and restore urban biodiversity, the need to study urban ecology has grown (Pierce et al., 2020). Unfortunately, ecological information on the effects of urbanization is often a limiting factor in designing and implementing effective biodiversity strategies (McDonnell, 2015; Shih et al., 2020). Improving urban native biodiversity requires developing ecological methods and interpretation of results that acknowledge the complexities of achieving biodiversity goals in urban areas (Pierce et al., 2020; Shwartz et al., 2014).

Amphibians are one of the most imperilled species assemblages with diversity and abundance declines reported globally (Hussain, 2012). Amphibians are considered a key indicator of ecological condition because of their low vagility and biphasic life cycle increasing their susceptibility to both aquatic and terrestrial stressors (Bridges & Semlitsch, 2000; Cushman, 2006; Sinsch, 1990). Urban environments pose a number of challenges to amphibian survival, such as habitat loss and fragmentation, direct mortality from road networks and built environments, and habitat degradation from pollutants and introduced predatory fish species (Calderon, 2019; Garcia-Gonzalez & Garcia-Vazquez, 2012; Konowalik et al., 2020; Rubbo & Kiesecker, 2005; Scheffers & Paszkowski, 2012).

Urban amphibian monitoring indicates a decreasing trend in amphibian diversity and abundance (Konowalik et al., 2020; Scheffers & Paszkowski, 2013). For example, a long-term monitoring programme in four cities in Poland reported decreasing numbers of amphibians at breeding sites, loss of breeding sites and complete disappearance of some species in recent decades (Konowalik et al., 2020). Urban amphibian studies report consistent patterns and trends, including substantial loss of pre-settlement natural wetlands, amphibian species richness and abundance decreases towards the city centre, dispersal routes compromised by road networks and wetland loss, and species-specific responses to urbanization (Hamer & McDonnell, 2008; Rubbo & Kiesecker, 2005). Despite numerous documented threats to amphibians from urbanization, and despite the loss of many amphibian species from cities, amphibians are still reported throughout the world in urban areas (Calderon, 2019; Konowalik et al., 2020; Scheffers & Paszkowski, 2013; Shulse et al., 2010; Simon et al., 2009; Skidds et al., 2007).

A review of conservation efforts for urban amphibians identified critical constraints such as limited lands available for conservation and a lack of ecological information available to inform and prioritize conservation (Scheffers & Paszkowski, 2012). Because urban conservation takes place in a complex and dynamic municipal decision-making environment, the need for clearly articulated, defensible, and socially supported conservation actions is acute (Shwartz et al., 2014; Turo & Gardiner, 2020). Translating ecological knowledge into conservation action requires the development of a conservation framework that identifies processes and tools needed to inform urban planning, management and restoration actions (Margules & Pressey, 2000).

We develop and test an urban conservation framework that can be used for any wildlife group, but here to support amphibian conservation (relating to urban planning and management activities) as a case study. Working with planners, residents and ecologists, we identified the eight critical steps in a framework for effective integration of conservation and management. Then we demonstrate the utility of the conservation framework using two amphibian species, wood frog and boreal chorus frog, which, although secure in Western Canada, have seen range declines within urban environments (B.C. Ministry of Environment, 2016; Hammerson, 2008). This adaptive approach facilitates easy application of our framework in other urban or non-urban landscapes with intense competition for land use.

2 MATERIALS AND METHODS

2.1 Urban amphibian conservation framework

To maintain amphibian species richness, associated viable populations and related ecological functions in a large urban centre, information on amphibian species in urban environments is needed for planning, management and restoration initiatives. To tackle this challenge, we formed an advisory committee with participation from academia, conservation organizations and municipal representatives. The panel included expertise in citizen science, wetland ecology, amphibian biology, landscape connectivity and municipal planning. Many municipalities are limited by staff time and expertise in their ability to conduct research and analyses needed for biodiversity conservation. These types of initiatives can be facilitated by forming collaborations among municipalities, academia and conservation organizations. The advisory committee met regularly over the duration of the programme to provide direction, feedback and establish process steps. These steps were developed adaptively and refined over time with input from an advisory committee established at the inception of an urban amphibian monitoring programme. This process arrived at an eight-step urban amphibian conservation framework.

2.2 Framework application

To highlight the utility of the urban amphibian conservation framework, we applied the framework in the city of Calgary, Alberta, Canada. A fundamental objective of Calgary's long-term plan is to maintain natural environments and ecological conditions able to support rich biodiversity and ecosystem services important to citizens (City of Calgary, 2004, 2015).

2.3 Study area

Calgary, Alberta, is one of Canada's largest cities, with a population of over 1.2 million. Typical of North American urban areas, Calgary has a heavily developed core surrounded by residential neighbourhoods that continue to spread, currently covering 848 km². As a result of this expansion, it is estimated that Calgary has lost 90% of its wetlands since European settlement began in the 18th century (City of Calgary, 2004) and most that remain in Calgary's urban environment contribute in some form to stormwater management. As of 2015, approximately 2729 wetlands remain within the city limits. These wetlands predominately occur on the edge of the city where densification has not yet occurred, alongside major roads within the transportation corridor, or within urban parks. Outside the city limits, the formerly predominate prairie pothole landscape remains somewhat intact but development and agricultural pressures occur in most neighbouring municipalities.

Historically, six amphibian species occurred within the current Calgary city limits: northern leopard frog (Lithobates pipiens), boreal chorus frog (Pseudacris maculata), wood frog (L. sylvaticus), Canadian toad (Anaxyrus hemiophrys), western toad (Anaxyrus boreas) and western tiger salamander (Ambystoma mavortium) (City of Calgary, 2014).

3 RESULTS

3.1 Urban amphibian conservation framework

Through facilitated meetings with an advisory group, we developed an eight-step amphibian conservation framework (Table 1). Steps 1–3 (goals, information, engagement) can occur in parallel and represent project scoping to outline conservation content, such as identifying the stakeholders, the programme goals and critical information needs (Schwartz et al., 2018). We present each step in two parts: the general description of the framework in an urban planning context and a specific description of how each step was conducted for amphibian conservation.

TABLE 1. Urban amphibian conservation framework

Step	Urban conservation issue	Research question	Methodological approach
1 Goals	Amphibian populations are at risk in urban areas.	What is the conservation goal for amphibians?	Clearly articulated conservation goals.
2 Information	Municipalities lack ecological information and/or planning practices that support coordinated citywide management and restoration decisions benefitting amphibians.	What information do decision-makers need to enable conservation action for amphibians?	Consultation with decision-makers and ecologists to identify existing information gaps and institutional barriers.
3 Engagement	Lack of public support for amphibian conservation.	How can we build social capital for investment in amphibian conservation?	Create opportunities for citizens to engage in knowledge production, learning and information sharing.
4 Monitoring	Limited amphibian monitoring data prevents inclusion based on amphibians in planning, management, and restoration.	What do we need to monitor to inform planning, management and restoration of amphibian habitat?	Development of a systematic monitoring programme.
5 Modelling	Urban landscape requires specific management recommendations of how amphibians respond to urban impacts.	What landscape and human features impact amphibian presence?	Occupancy modelling
6 Habitat ID	Investment for wetland protection and restoration is limited and prioritization is required.	Where is priority amphibian habitat?	Habitat modelling to identify high-valued amphibian habitat.
7 Connectivity	Urban landscapes present many dispersal barriers to amphibians.	Where are the amphibian dispersal pathways and where are barriers?	Connectivity modelling to determine potential amphibian movement pathways.
8 Integration	Conservation requires the implementation of ecological findings into local area and citywide planning processes, management and restoration plans, policies and decision tools.	How do we integrate results into existing planning, management and restoration approaches?	Identify existing planning processes, management and restoration plans, policy and tools that results can inform. Identify opportunities for integration of results.

3.1.1 Step 1: Goals

Framework. An important component of project scoping is to articulate conservation goals and objectives. Programme collaborators were asked to identify target species, assess common threats, identify critical information needs and outline the overall conversation goal. The response of species to urbanization is an important consideration in defining the conservation goal (Hamer & McDonnell, 2008; Hamer & Parris, 2011).

Urban conservation goals may focus on maintaining existing populations, improving species richness or restoring populations extirpated from the urban environment. Conservation goals can be iteratively developed through steps 2 and 3, where collaborators are engaged to ensure the social, cultural and political contexts are considered.

Amphibian conservation. The advisory committee identified three objectives to support an agreed upon conservation goal: address ecological knowledge gaps in amphibian ecology, build social capital and implement findings into planning, management, and restoration plans and tools.

3.1.2 Step 2: Information

Framework. It is important to consult with decision-makers and ecologists to identify information and institutional process gaps. What ecological information do we need to have to inform biodiversity planning, management, and policy? These questions are best answered by consulting with City representatives and subject matter experts early in the process. Cities tend to plan at local development unit scales, fragmenting conserving biodiversity goals at a landscape level, potentially leading to what Allred et al. (2021) described as a series of asynchronous or ‘uncoordinated, local planning decisions’. Consulting with City representatives early ensures that the final step in any process (research results and products integrated into the key policies, strategic plans and management operations) is not the first time local policy-makers are engaged (Allred et al., 2021; Sterling et al., 2017).

Amphibian conservation. City of Calgary representatives and amphibian experts identified numerous technical knowledge gaps reducing the City's ability to plan and manage for amphibians. These included an inventory of amphibian species, where they occur, how amphibians are responding to urbanization, and an amphibian movement network to support dispersal among wetland habitats. The process of prioritization of areas for habitat and dispersal pathway protection and restoration was deemed an important gap to fill as the built footprint of the City continues to expand, removing wetland and upland amphibian habitat.

3.1.3 Step 3: Engagement

Framework. Creating opportunities for the public to engage in knowledge production, learning and sharing is an important component to fostering voices to support necessary biodiversity conservation. An engaged public is essential as municipal governments consider how to allocate limited funds to the many competing public interests potentially resulting in public support for investment in biodiversity conservation. This was demonstrated in New York's Hudson Valley where external political pressure increased political support for biodiversity action (Allred et al., 2021). Therefore, building public support for amphibian conversation is an important component of urban conservation success (Schwartz, 2006).

Citizen science (also referred to as community science with the understanding that one does need not be a citizen to be valued) is a useful approach for addressing data gaps while also engaging the public in research and conservation (Cooper et al., 2021; Marsh & Cosentino, 2019; McKinley et al., 2015; Westgate et al., 2015). Research suggests that actively involving the public in science can improve biological knowledge, conservation awareness and encourage participation in decision-making processes (Aceves-Bueno et al., 2015; Shirk et al., 2012).

Amphibian conservation. The opportunities programme developed a 3-year citizen science initiative to help monitor amphibians using call surveys and physical observations. Important components of the initiative included messaging about the need for conservation of wetlands and amphibians and providing direction on how the public can participate in city planning decisions.

The citizen science platform resulted in civic engagement to support amphibian conservation for 546 Calgarians with many contributing to data collection (approximately 100 per year) or attending a programme sponsored event. Project and modelling results were shared with participants via infographics and story boards. Lee et al. (2021) outline the benefits of engagement to urban conservation and provide recommendations for citizen science programme design based on lessons learned.

3.1.4 Step 4: Monitoring

Framework. Development of a systematic wildlife monitoring programme is the platform for generating ecological information to inform and update conservation decision-making. In particular, animals can be difficult to detect given diverse habitats, diurnal/seasonal movements among habitats, small size of many urban wildlife species and morphologically distinct life forms (Muths et al., 2005). There are numerous pre-existing methods, such as autonomous recording units (ARUs), call surveys, DNA for monitoring and analytics to identify and monitor wildlife in urban settings (Garcia-Gonzalez & Garcia-Vazquez, 2012; Lankau, 2015; Shulse et al., 2010). Determining the best method depends on overall programme goals, resources available and geographic location of the study.

Amphibian conservation. The amphibian monitoring programme systematically monitored wetlands after dusk using call surveys and during the day using physical observations. Monitoring data were combined with ARUs placed at a small number of sites as a quality control measure (Lee et al., 2021).

We randomly identified 250 wetland sites, and, after field visits to determine safety and ease of public access and landownership, selected 42 survey sites. To accommodate permitting and the need for public accessibility, sites were located on municipal or provincial crown land. We also favoured permanent wetland sites as they were easier for the public to locate and were more common in the urban environment.

Survey methods were based on the provincial Sensitive Species Inventory Guidelines (Government of Alberta, 2013) and included visual and call surveys. We recruited members of the public to conduct nine systematic surveys of each wetland site per season to determine amphibian presence throughout the amphibian active season (April to August) from 2017 to 2019. The resulting 3-year amphibian dataset was used as the basis for occupancy and informed connectivity modelling.

After 3 years of amphibian monitoring at study wetlands, we documented three of the six amphibian species historically reported through provincial records. We did not observe northern leopard frog, Canadian toad, or western toad, but did detect boreal chorus frog, wood frog and western tiger salamander.

Boreal chorus frog was the most common species, with observations reported at 36 sites (85% of sites). Wood frogs were observed at 15 (36% of sites). Twelve sites (29%) reported co-occurrence of the two species. Only six sites had no amphibian observations during the 3-year study. Although we generalized data to amphibian presence for modelling, evidence of breeding (eggs and/or tadpoles) occurred at 12 of the sites representing 29% of the urban wetland sites surveyed. Amphibian observation and wetland survey sites can be downloaded from Zenodo: https://doi.org/10.5281/zenodo.6251519 (Sanderson, 2022a).

3.1.5 Steps 5 and 6: Modelling and habitat ID

Framework. Urban wildlife lives in a built environment make it important to understand how they respond to artificial landscape elements (Browne et al., 2009a; Shulse et al., 2010). Occupancy modelling (step 5) can be used to explore space use by wildlife in relation to landscape features while accounting for imperfect detection (MacKenzie et al., 2017). In combination, habitat suitability modelling (step 6) can identify high-quality habitat in the landscape based on an individual response to sets of environmental cues (Milanovich et al., 2012; Mushet et al., 2012).

Amphibian conservation. To identify highly valued habitat and probable movement pathways, we outlined an analytical process that used occupancy modelling (step 5) to parameterize habitat suitability models, identify priority wetlands (step 6) and develop connectivity models (step 7) (Figure 1). Modelling was completed by technical experts at universities/NGOs with the consultation of civic and other partners.

Details are in the caption following the image — **FIGURE 1**
Open in figure viewer PowerPoint

Analytical process for identification of high priority wetlands and movement pathways

Modelling. We used multi-season occupancy models to explore amphibian occupancy patterns for which there were sufficient observations (i.e., wood frogs and boreal chorus frogs) while accounting for imperfect detection (MacKenzie et al., 2017). Western tiger salamander was not included in the modelling as there were insufficient observations. We binned data into five monthly periods (April, May, June, July and August) given that previous analyses have shown that the probability of detection varied by month for boreal chorus frogs and varied linearly with month for wood frogs. To reduce the bias of occupancy estimates, we constructed models that allowed the probability of detection to vary by month for boreal chorus frogs and linearly by month for wood frogs. We held colonization (γ) constant as this was not a parameter of interest, fit single landcover covariates on occupancy (ψ), and compared these results to the null model (Randall et al., 2015).

We normalized proportional data using a logit transformation (Warton & Hui, 2011) and standardized all remaining variables. We tested if landscape variables were correlated and, if they were, we kept the top-ranked model variable and eliminated the others from the final model set. Environmental conditions, such as water quality, were not included in our surveys so we were unable to determine how pollution or fish predation may be impacting amphibians. We used Akaike's information criterion (AIC) to select the most parsimonious model and calculate a model weight (Burnham & Anderson, 2002) and only considered models that were ranked above or within Δ2 AIC of the null model as important for predicting amphibian occupancy. Other models were excluded from the results and were not used to construct the habitat suitability index (HSI).

We generated landscape variables from GIS data available on the City of Calgary's open web portal to use in occupancy modelling (Table S1 in the Supporting Information). Data can be downloaded from the City of Calgary open data portal: https://data.calgary.ca/ (City of Calgary, 2021).

The top-ranked model for wood frog, with 97% of the AIC weight, showed that wood frog occurrence increased as distance to forest decreased (Table 2). Wood frog occurrence also increased with proportion of grasslands in the 20 m buffer (< 2% AIC weight), proportion of water in the 50 m buffer (< 1%), and with wetland size (< 1%). Wood frog occurrence declined with the percentage of impervious surfaces (<1%), with slope (<1%), and as distance to roads decreased (<1%).

TABLE 2. Wood frog (Lithobates sylvaticus) occupancy in Calgary, Alberta, Canada, in relation to landscape variables. Occupancy determined from results of call surveys conducted from April to August (2017–2019)

Model	−2 LL	K	AIC	ΔAIC	w_i
Ψ (Distance to forest)	145.86	5	155.86	0.00	0.97
Ψ (Proportion of grassland in 20 m buffer)	153.89	5	163.89	8.03	0.02
Ψ (Proportion impervious surface)	157.46	5	167.46	11.60	<0.01
Ψ (Distance to nearest road)	157.55	5	167.55	11.69	<0.01
Ψ (Null)	160.87	4	168.87	13.01	<0.01
Ψ (Proportion of water in 50 m buffer)	159.88	5	169.88	14.02	<0.01
Ψ (Slope within 50 m buffer)	160.35	5	170.35	14.49	<0.01
Ψ (Size of wetland)	160.75	5	170.75	14.89	<0.01

Abbreviations: −2 LL , −2 log likelihood; AIC, Akaike information criterion; K, the number of parameters; w_i , Akaike weight; ψ, occupancy.

The top-ranked model for boreal chorus frog, with 60% of the AIC weight, showed that their occurrence increased with the proportion of manicured landscape (Table 3). Boreal chorus frog occurrence declined with proportion of water (21% AIC weight), slope (4%), proportion of forest (4%), proportion impervious surface (3%) and as distance to nearest wetland increased (2%). Occurrence increased as the proportion of grasslands increased (3%).

TABLE 3. Boreal chorus frog (Pseudacris maculata) occupancy in Calgary, Alberta, Canada, in relation to landscape variables. Occupancy determined from results of call surveys conducted from April to August (2017–2019)

Model	−2 LL	K	AIC	ΔAIC	w_i
Ψ (Proportion of manicured in 20 m buffer)	401.38	8	417.38	0	0.60
Ψ (Proportion of water in 100 m buffer)	403.51	8	419.51	2.13	0.21
Ψ (Null)	408.97	7	422.97	5.59	0.04
Ψ (Slope within 50 m buffer)	407.01	8	423.01	5.63	0.04
Ψ (Proportion of forest in 20 m buffer)	407.06	8	423.06	5.68	0.04
Ψ (Proportion impervious surfaces in 20 m buffer)	407.44	8	423.44	6.06	0.03
Ψ (Proportion grassland in 100 m buffer)	401.38	8	423.49	6.11	0.03
Ψ (Distance to nearest wetland)	403.51	8	424.02	6.64	0.02

Abbreviations: −2 LL, −2 log likelihood; AIC, Akaike information criterion; K, the number of parameters; w_i, Akaike weight; ψ = occupancy.

Habitat ID. We used variables identified in occupancy models to construct an HSI for wood frog and boreal chorus frog. Wetlands were buffered by 250 m for wood frog and 100 m for boreal chorus frog to represent upland habitat and the area required to support seasonal movements (Baldwin et al., 2006; Scherer et al., 2012). The AIC weights generated from occupancy modelling were used to adjust the importance of each covariate in HSI models. Distribution curves derived from the occupancy models were used to determine suitability values for each covariate in the HSI model. GIS covariates were then combined using “Map Algebra” formulas to determine potential site suitability.

HSI values range from 0 to 1, where 1 represents the high valued habitat. We calculated the mean HSI value for each wetland and its buffer for both amphibian species and discretized this value into three classes to represent low (0–0.3), medium (0.3–0.6), and high (>0.6) valued habitat based on expert opinion. We validated HSI models using opportunistic observations (observed at non-survey wetlands) reported during the 3-year amphibian monitoring programme, including 68 boreal chorus frog and 17 wood frog observations. High-valued habitats for each species were overlain. Areas of high overlap represent priority areas of habitat for amphibians occurring in Calgary under current conditions.

To identify the most suitable habitat for wood frogs, we used the four attributes with the largest impact on wood frog occurrence: distance to forest, proportion of grassland, proportion of impervious surface and distance to roads. For boreal chorus frogs, we used five attributes that had the largest impact on boreal chorus frog occurrence: proportion of manicured land cover, slopes, proportion of forest, proportion of impervious surface and proportion of grassland.

To derive HSI models we used weightings (AICW) from the occupancy models to derive the formulas:

\begin{eqnarray} && {\rm{HSI}}\;{\rm{wood}}\;{\rm{frog}}\ =\ (\rm{Distance\ to\ forest} \times {\rm{0.97}}) + (\rm{Grassland} \times {\rm{0.02}}) \nonumber\\ &&\quad +\, (\rm{Impervious\ surface} \times {\rm{0.005}}) + (\rm{Distance\ to\ roads \times} {\rm{0.005})}. \end{eqnarray}

(1)

\begin{eqnarray} &&\hspace*{-9pt} {\rm{HSI}}\;{\rm{boreal\ chorus\ frog}} = (\rm{Manicured} \times {\rm{0.82}}) + (\rm{Slope} \times {\rm{0.05}}) \nonumber\\ && +\, (\rm{Forest} \times {\rm{0.05}}) + (\rm{Impervious} \times {\rm{0.04}}) + {\rm{Grassland}} \times {\rm{0.04}}). \end{eqnarray}

(2)

We mapped high-quality habitats for wood frog and boreal chorus frog to validate our observation data (Figure 2). For model validation, we used opportunistic observations (not at specified survey sites) reported by citizens during the amphibian monitoring programme. There were 17 for wood frog and 68 for boreal chorus frog observations reported by citizens. 71% of the wood frog and 58% of boreal chorus frog opportunistic observations occurred in the high habitat value from the HSI model.

3.1.6 Step 7: Connectivity

Framework. A key threat to urban wildlife populations is habitat fragmentation reducing functional connectivity between habitat and resource patches (Hamer et al., 2012). Functional connectivity is the ability of a species to move between habitat and resource patches.

A common approach for identifying wildlife movement pathways is to use connectivity software to model landscape permeability (Churko, 2016; Nowakowski et al., 2015). Koen et al. (2014) developed a method to identify areas of high landscape connectivity for multiple (amphibian) species based on landscape structure. They created a cost surface to represent landscape permeability and used circuit theory to connect randomly generated focal nodes (McRae et al., 2008). If possible, it is important to develop connectivity models for each species of concern to account for interspecific differences in preferred habitat and dispersal distances (Browne et al., 2009b; Jarchow et al., 2016).

Amphibian conservation. Amphibians rely on a breeding pond network, among which juveniles are able to disperse and move to new habitats (Cushman, 2006). This is a series of connected habitats that are permeable to amphibian movement, including ephemeral wetlands, that can increase the effective dispersal distance between ponds (Allen et al., 2020). Amphibians are prone to local extinction events and depend on recolonization from other breeding ponds for survival (Cushman, 2006). Zamberletti et al. (2018) studied the degree of wetland connectivity and found management actions (protection and restoration) for a specific wetland should depend on its location within the broader landscape, highlighting the importance of conservation action focused on wetland networks and not isolated breeding ponds. In urban areas, where road networks and the built environment fragment habitats, options for amphibian movement are greatly reduced resulting in low functional connectivity and reduced population resilience due to isolation, increased risk of local extinction and/or inbreeding (Allen et al., 2020; Koen et al., 2014; Zaffaroni et al., 2019; Zamberletti et al., 2018).

We developed functional connectivity models for two amphibian species based on electrical circuit theory and then summed the models to identify intersections between them as a step towards prioritizing areas for urban conservation (Boyle et al., 2017; Koen et al., 2014). We developed landscape resistance surfaces based on expert knowledge to represent landscape permeability and then ran the current among focal nodes of high-quality habitats generated from occupancy and habitat suitability models in the initial steps to identify potential movement pathways (Cushman & Landguth, 2012).

We used Circuitscape software and inputs of a resistance surface and focal nodes to identify probable movement pathways for amphibians (McRae et al., 2008). The study area was buffered by 20% to account for edge effects in modelling (Koen et al., 2014). Focal nodes were generated based on high valued habitat identified in the HSI models. An expert-based approach was used to create resistance surfaces for amphibian species to identify potential amphibian movement pathways within the city limits. Landscape attributes were selected by experts for each amphibian species observed in Calgary during the 3-year monitoring period (i.e. wood frog, boreal chorus frog and western tiger salamander) and habitat connectivity modelled only for the two frog species, as there were too few western tiger salamander occurrences reported during the study. Each landscape attribute was classified into categories for resistance (i.e. not permeable) to amphibians (Table 4). Each feature was given a resistance class of habitat, favourable matrix, less favourable matrix or strong barrier (Churko, 2016) (Table S2 in the Supporting Information). Resistance classes were converted to numerical values based on three resistance scenarios: sigmoidal, logarithmic and exponential (Table 5). We applied a cut-off of 1000 m for wood frog and 600 m for boreal chorus frog to account for maximum dispersal distances (Berven & Thaddeus, 2010; Bishir et al., 2018; Dodd, 2013).

TABLE 4. Landscape variables and classifications used to build amphibian resistance surfaces

Transportation	Hydrology	Landcover	Impervious surface	Slope
>4-lane paved roads	River	Forest	Buildings	>20°
Four-lane paved roads	Creek	Grassland	Gravel patches	16–20°
Neighbourhood roads	Canal	Shrubland	pavement or concrete	12–15°
LRT track	Reservoir	Agriculture crop		0–11°
Laneways	Wetland	Agriculture pasture
park pathway		Golf course
railway		Manicured grass
		Sports facility
		Bare ground
		Construction

TABLE 5. Conversion of resistance surface classes into numerical values

Resistance value	Sigmoidal	Exponential	Linear
Habitat	1	1	1
Favourable matrix	100	10	333
Less favourable matrix	900	100	666
Strong barrier	1000	1000	1000

We used Circuitscape (Version 5.0) to run connectivity models for both species on each resistance scenario and focal node placement (Anantharaman et al., 2019). These models predict probabilistic movement along all possible paths and assume animals do not know the landscape (Wade et al., 2015). The three resistance scenarios for each species were summed to generate a current density map for each species, and we took the top 50% of the summed resistance model for each species and created a binary surface. These surfaces were overlain to identify movement pathways with areas of high overlap indicating the most likely wetlands and upland habitat for amphibian movement in Calgary under current conditions (Figure 3).

To better inform planning, we used modelling results to identify core wetlands (wetland sites supporting high valued habitat for both amphibian species in Calgary) and potential movement pathways for amphibians (top 50% of summed species connectivity models; Figure 4a). A key consideration in conservation planning and management is land ownership as conservation strategies and implementation could depend on who has jurisdiction. In Figure 4b, we highlight government-owned land (municipal and provincial), where some level of protection already occurs or a government agency has control over the management of wetland and upland habitat. Conservation strategies and implementation could depend on who has jurisdiction. An analysis of potential barriers to amphibian movement identified roads as a key barrier, demonstrating a high level of fragmentation in the potential network (Figure S1 in the Supporting Information). Modelling datasets can be downloaded from Zenodo: https://doi.org/10.5281/zenodo.6233027 (Sanderson, 2022b).

3.1.7 Step 8: Integration

Framework. Ecological information generated from the urban conservation framework can be integrated into urban planning, park and water management and restoration projects to support urban biodiversity. Engaging decision-makers early (step 2) will help to identify where models and results need to be integrated to ensure consideration in planning, management and restoration projects into existing plans, policies and tools. In urban areas, where conservation dollars may be limited, mitigation expensive, and land-use pressure significant, conservation practitioners may choose to focus on proactively managing for amphibian species that are doing well in the urban landscape (Sterrett et al., 2019). This step can also be used to identify limitations in city planning, policy, or management that would need to be addressed to maintain wildlife populations as well as support habitat networks. For example, framework results could identify if stormwater ponds constructed for the purpose of controlling water quality and quantity in urban areas are contributing to aquatic habitat. Studies indicate artificially constructed ponds can be designed to support aquatic habitats, and there are numerous design considerations that could be implemented if stormwater ponds are not currently supporting wildlife populations (Clevenot et al., 2018; Holtmann et al., 2017).

Amphibian conservation. Decision-makers identified two key City of Calgary departments (Water Management and Parks), seven plans (e.g. biodiversity strategy) and an internal decision support tool (restoration prioritization tool) that would benefit from the results of the framework. Amphibian modelling spatial datasets and accompanying technical reporting were shared with both City of Calgary Water Management and Urban Parks Departments.

The information is now being considered by City planners to inform policy (e.g. identify ecologically sensitive areas for protection as environmental reserves, including wetland/amphibian connectivity in the development of master drainage plans), neighbourhood planning (e.g. inform area structure plans (subdivision planning) within the development process), and strategic planning (e.g. support an ecological network as outlined in municipal development plan) objectives for citywide biodiversity connections.

To get the most out of limited conservation resources, high-priority wetlands and amphibian movement pathways can lead to improved management and restoration actions. For example, results were used to inform parks ecologists of the presence of amphibian species resulting in the use of beneficial management practices during a restoration project (e.g. timing of plantings to avoid migration of amphibian species to and from water, clumping of vegetation to enhance amphibian cover). Future considerations include incorporating model results into existing restoration prioritization tools.

The success of the framework will ultimately be measured by how ecological knowledge is integrated into planning and management at the municipal level. Continued monitoring over time will determine amphibian responses, positive or negative, to conservation actions. Ideally, a timeframe could be assigned to evaluate the success of amphibian conservation resulting from the implementation of the framework.

4 DISCUSSION

Amphibian conservation in urban areas is challenged by development pressure, competing for land uses, limited funding and a lack of ecological knowledge (i.e. species diversity, core amphibian habitat and amphibian movement pathways) to support citywide planning and management initiatives (McDonnell, 2015; Pierce et al., 2020; Shih et al., 2020). These challenges can prevent the implementation of effective conservation actions in support of biodiversity in urban areas (Sandström et al., 2006). A recent global survey of urban experts identified lack of technical knowledge as a key barrier to biodiversity conservation and emphasized the need for frameworks and tools to support urban conservation (Shih et al., 2020). We developed an urban conservation framework as a process to adaptively manage wildlife richness, associated viable populations and related ecological functions in a large urban area. We demonstrated the utility of this framework through an application in Calgary, Alberta, Canada, for amphibian conservation.

4.1 Insights gained through framework application

Using a structured process and well-established methods allowed us to provide ecological information to decision-makers that can guide policy, planning and management decisions related to amphibian conservation. Along the way, we identified insights that are broadly applicable to other conservation practitioners. The importance of early engagement of decision-makers cannot be overemphasized and the ability to identify critical information gaps, engage the public, document who makes decisions and show how information can be integrated into policy and planning were all key and interrelated steps (Schwartz et al., 2018).

Decision-makers play an important role in identifying knowledge gaps relevant to planning and policies and identifying products useful in the decision-making process. A recent study that surveyed urban planners about biodiversity concluded a lack of technical knowledge and resources reduced their ability to plan for biodiversity (Sandström et al., 2006). Urban planners noted that legislation was an important driver in biodiversity planning. Early engagement of decision-makers from appropriate departments at the City of Calgary resulted in identifying useful ecological knowledge, in our case for amphibians, and identified existing plans and policies where amphibian biodiversity information could be incorporated (City of Calgary, 1994, 2015, 2017).

To fill knowledge gaps, we blended empirical data collected by the public and expert knowledge. By engaging citizen scientists in data collection, we were able to communicate the importance of wetlands and amphibian conservation in ways more effective than through the media or scientific reports (McKinley et al., 2017). Research indicates the importance of building social capital to support biodiversity conservation (Schwartz, 2006). For example, citizen science programmes have shown that engaged members of the community can be a strong voice for conservation to influence city policy and planners (Bonney et al., 2021). We highlight the value of a citizen science approach, such as spreading of conservation messaging through participants to the broader community and increasing interest in participating in civic planning processes (Lee et al., 2021).

In an urban environment, conservation actions often need to consider multi-jurisdictional land ownership and operational management that increase the number and type of conservation actions required. For example, stormwater ponds have been shown to support amphibian biodiversity in large urban centres, but management has historically focused on their function in storing stormwater not on promoting biodiversity (Hamer & Parris, 2011; Scheffers & Paszkowski, 2012). Efforts to promote amphibian conservation outside of a natural area may require coordination with departments responsible for management whose purposes are not maintaining biodiversity. Our process highlighted the importance of cross-department engagement. For example, staff working on local area plans were provided with access to models of high valued amphibian habitat and movement pathways to encourage retention of wetlands and existing potential movement pathways. In addition, the City of Calgary is developing a master drainage plan, and the amphibian models and modelling results were provided to the city planners to encourage retention of core wetlands and movement pathways.

4.2 Framework application in Calgary

The urban amphibian conservation framework was used to provide City of Calgary decision-makers with specific ecological knowledge for conservation-oriented planning and management decisions. Prior to this study, little was known about which amphibian species were present within the City of Calgary, where they were located, habitat associations and movement barriers.

Of the six species that historically occupied the City of Calgary, only three remain: boreal chorus frog, wood frog and western tiger salamander. Of these species, only boreal chorus frogs are commonly found at Calgary's wetlands. This is likely because the boreal chorus frog is a generalist species that occupies a wide range of wetlands of different sizes, types and permanency (Dodd, 2013; Rubbo & Kiesecker, 2005). Boreal chorus frog occupancy increased with the amount of manicured (mowed) grassy areas, which may not be surprising considering it is a grassland prairie species on the Great Plains and tends to prefer short grassy vegetation (Dodd, 2013). Perhaps more surprising was the negative association with the proportion of water. Boreal chorus frogs are often associated with small, ephemeral wetlands (Dodd, 2013); however, the wetlands in our study area tended to be larger permanent wetlands and the total water landcover layer included landscape features such as rivers and reservoirs that were likely not suitable habitat for boreal chorus frogs. However, boreal chorus frog occurrence did increase when other wetlands were near as we predicted.

Unsurprisingly, areas identified as high-valued habitat and with good connectivity were typically located on the outskirts of Calgary where there has been less development and fewer wetlands drained. Given the strong association with forested areas, wood frogs were more– commonly associated with wetlands in areas with the intact forest that are also more common at the city outskirts (Rubbo & Kiesecker, 2005). Protecting intact wetlands is far more cost and time effective than complete restoration. This information is also valuable to support city planning teams to both identify high-valued habitat for protection at the scale of the area structure plan (for urban subdivision) within the development process and to achieve strategic planning objectives for citywide biodiversity connections.

Although most high-valued habitat was located on the edge of Calgary, we also identified some within the City core. But these generally had low connectivity. Amphibians are typically distributed as metapopulations, with individual populations experiencing periodic extinction and re-establishment through colonization requiring movement among ponds (Smith & Green, 2005). Amphibian populations in the core of the City are at risk of extirpation due to stochastic events, such as pond drying or flooding. Due to the distance among sub-populations and fragmentation, there is limited recolonization potential (Parris, 2006). There are two strategies to re-establish extirpated populations within the core: restore connectivity or conservation translocation. Connectivity could be improved through mitigation measures such as restoring natural dispersal pathways like creeks and streams, or by building crossing structures that span specific barriers. These latter mitigations have shown some utility in the urban environment (see conservation evidence) but are often expensive and require ongoing maintenance (Beebee, 2013; Helldin & Petrovan, 2019). Alternatively, conservation translocations have been widely used for restoring amphibian populations but rarely in the urban environment (Linhoff et al., 2021; Randall et al., 2018; Soorae, 2016).

4.3 Further considerations for urban amphibian conservation

Measuring success of the framework to support amphibian conservation in urban centres requires continued monitoring to track amphibian species richness, viable populations and environmental function over time. We recommend a renewed focus on identification of breeding ponds, as this was complicated by public participation and challenges in identifying eggs and tadpoles to the species level. We recommend expanding monitoring to measure environmental conditions that affect the survival of amphibian populations to support development of beneficial management practices. Additionally, we did not have knowledge of culvert locations which could act as conduits for amphibian movement across roads. We recommend further investigation into road sections identified as barriers in connectivity modelling to assess if existing infrastructure could be modified to support amphibian movement.

A challenge to implementing urban conservation strategies for amphibians is a lack of clearly articulated population targets or ecological decision-making triggers to drive conservation initiatives such as habitat restoration or road mitigations to enable dispersal. As outlined by Shih et al. (2020), policy triggers for implementing conservation actions in urban areas are important. Our urban conservation amphibian framework provides the ecological knowledge necessary to know where protection, conservation or restoration action is needed, but without strong policy direction, implementation will be challenged.

4.4 Broader application

It is important to recognize that no framework can address all conservation planning and decision-making challenges; there are diverse tools (i.e. spatial modelling, cost assessments, scenario planning) available to conservation practitioners (Schwartz, 2006). Conservation frameworks are widespread and have been developed to address regional protected areas planning (Groves et al., 2002), provide strategic foresight to predict future conditions (Cook et al., 2014) and identify where best to deploy specific conservation actions (Sarkar et al., 2006). Conservation practitioners need to clearly define objectives, evaluate missing critical information and understand the decision-making environment prior to selecting an appropriate framework for conservation. In our urban amphibian conservation framework, steps 1–3 addressed these questions and enabled conservation practitioners to select appropriate conservation tools.

By implementing the urban amphibian conservation framework, we improved ecological information using well-established methods in amphibian conservation (steps 4–7) to enable better planning, management and restoration decisions. Furthermore, we demonstrated the importance of early identification of existing municipal plans, policy and tools to improve conservation action.

Amphibian species were selected as an ecological indicator because of their low vagility and biphasic life cycle increasing their susceptibility to both aquatic and terrestrial stressors which are exacerbated in the urban environment (Bridges & Semlitsch, 2000; Cushman, 2006; Sinsch, 1990). To maintain amphibians in the urban environment requires a concerted effort in conservation planning and management. Our framework is widely applicable to other large urban centres or ex-urban landscapes with intense competition for land use.

ACKNOWLEDGEMENTS

Amphibian monitoring data for this study was generated by the Call of the Wetland Program, which was generously funded by Alberta Innovates, Enbridge, Calgary Foundation, Alberta Ecotrust Foundation, RBC Tech for Nature Grant via The City of Calgary, Bow River Basin Council, TD Friends of the Environment, World Wildlife Fund and Mount Royal University Institute for Environmental Sustainability. In addition, we thank all the Calgarians who participated in data collection via the Call of the Wetland Program. Thank you to Natalie Lora Colquhoun, Jen Demone, Rachel Pizante and Coral Sawatzky for supporting the project during a summer internship. Thank you to Dr. John Wilmshurst for reviewing the manuscript to assist with structure and copy-edit.

CONFLICT OF INTEREST

There are no known conflicts of interest.

AUTHOR CONTRIBUTIONS

T.S.L., D.D. and L.A.R. conceived the ideas and designed the methodology, with additional input by V.A.C., H.R. and I.F.C.; N.L.K. and T.M.B. coordinated the citizen scientists; K.S. and N.L.K. collated and prepared the data; T.S.L. and L.A.R. analysed the data and drafted the manuscript, with critical revisions by D.D., N.L.K., T.M.B., H.L.K. and A.M. All authors contributed to the drafts and gave final approval for publication.

Open Research

PEER REVIEW

The peer review history for this article is available at https://publons.com/publon/10.1002/2688-8319.12139

DATA AVAILABILITY STATEMENT

Amphibian species occurrence used in this analysis was generated by Call of the Wetland Amphibian Monitoring Program led by Miistakis Institute, raw monitoring data can be downloaded at zenodo: https://doi.org/10.5281/zenodo.6251519 (Sanderson, 2022a). Amphibian modelling results can be downloaded at zenodo: https://doi.org/10.5281/zenodo.6233027 (Sanderson, 2022b). Datasets used in the analysis (Table S1 in the Supporting Information) are available on the City of Calgary open data portal: https://data.calgary.ca/ (City of Calgary, 2021). From the open data portal, we used land cover data (includes grasslands, forests, manicured grass, water bodies (including wetlands), and built-up areas: https://data.calgary.ca/Environment/Citywide-Land-Cover/as2i-6z3n, and major road network: https://data.calgary.ca/Transportation-Transit/Major-Road-Network/tqjs-vnhy.

Supporting Information

REFERENCES

Aceves-Bueno, E., Adeleye, A. S., Bradley, D., Brandt, W. T., Callery, P., Feraud, M., Garner, K. L., Gentry, R., Huang, Y., McCullough, I., Pearlman, I., Sutherland, S. A., Wilkinson, W., Yang, Y., Zink, T., Anderson, S. E., & Tague, C. (2015). Citizen science as an approach for overcoming insufficient monitoring and inadequate stakeholder buy-in in adaptive management: Criteria and evidence. Ecosystems, 18, 493–506. https://doi.org/10.1007/s10021-015-9842-4
10.1007/s10021‐015‐9842‐4
Web of Science®Google Scholar
Allen, C., Gonzales, R., & Parrott, L. (2020). Modelling the contribution of ephemeral wetlands to landscape connectivity. Ecological Modelling, 419, 108944. https://doi.org/10.1016/j.ecolmodel.2020.108944
10.1016/j.ecolmodel.2020.108944
Web of Science®Google Scholar
Allred, S., Stedman, R., Heady, L., & Strong, K. (2021). Incorporating biodiversity in municipal land-use planning: An assessment of technical assistance, policy capacity, and conservation outcomes in New York's Hudson Valley. Land Use Policy, 104, 1–17. https://doi.org/10.1016/j.landusepol.2021.105344
10.1016/j.landusepol.2021.105344
Web of Science®Google Scholar
Anantharaman, R., Kimberly, H., Viral, S., & Lan, E. (2019). Circuitscape in Julia: High performance connectivity modelling to support conservation decisions [Preprint]. arXiv 1906.03542.
Google Scholar
Baldwin, R. F., Calhoun, A. J. K., & deMaynadier, P. G. (2006). Conservation planning for amphibian species with complex habitat requirements: A case study using movements and habitat selection of the Wood Frog Rana sylvatica. Journal of Herpetology, 40(4), 442–453. https://doi.org/10.1670/0022-1511(2006)40[442:cpfasw]2.0.co;2
10.1670/0022?1511(2006)40
Web of Science®Google Scholar
B.C. Ministry of Environment. (2016). Conservation status report: Lithobates sylvaticus. B.C. Conservation Data Centre. https://a100.gov.bc.ca/pub/eswp/esr.do;jsessionid=XZvvhqrCG6xH7J4X1wswvxb7Dpv5BFglppl3GCQ28kTvdGTTFqJG!-378308228?id=14253
Google Scholar
Beebee, T. J. C. (2013). Effects of road mortality and mitigation measures on amphibian populations. Conservation Biology, 27, 657–668. https://doi.org/10.1111/cobi.12063
10.1111/cobi.12063
PubMedWeb of Science®Google Scholar
Berven, K. A., & Thaddeus, A. (2010). Dispersal in the wood frog (Rana sylvatica): Implications for genetic population structure. Society for the Study of Evolution Stable, 44(8), 2047–2056. https://doi.org/10.1111/j.1558-5646.1990.tb04310.x
10.1111/j.1558?5646.1990.tb04310.x
Google Scholar
Bishir, S. C., Hossack, B. R., Fishback, L. A., & Davenport, J. M. (2018). Post-breeding movement and habitat use by wood frogs along an Arctic–Subarctic ecotone. Arctic, Antarctic, and Alpine Research, 50(1),. https://doi.org/10.1080/15230430.2018.1487657
10.1080/15230430.2018.1487657
Web of Science®Google Scholar
Bonney, R., Byrd, J., Carmichael, J. T., Cunningham, L., Oremland, L., Shirk, J., & Von Harten, A. (2021). Sea change: Using citizen science to inform fisheries management. Bioscience, 71(5), 519–530. https://doi.org/10.1093/biosci/biab016
10.1093/biosci/biab016
PubMedWeb of Science®Google Scholar
Boyle, S. P., Litzgus, J. D., & Lesbarrères, D. (2017). Comparison of road surveys and circuit theory to predict hotspot locations for implementing road-effect mitigation. Biodiversity and Conservation, 26(14), 3445–3463. https://doi.org/10.1007/s10531-017-1414-9
10.1007/s10531?017?1414?9
Web of Science®Google Scholar
Bridges, C. M., & Semlitsch, R. D. (2000). Variation in pesticide tolerance of tadpoles among and within species of Ranidae and patterns of amphibian decline. Conservation Biology, 14, 1490–1499. https://doi.org/10.1046/j.1523-1739.2000.99343.x.
10.1046/j.1523‐1739.2000.99343.x
Web of Science®Google Scholar
Browne, C. L., Paszkowski, C. A., Foote, A. L., Moenting, A., & Boss, S. M. (2009a). The relationship of amphibian abundance to habitat features across spatial scales in the Boreal Plains. Écoscience, 16(2), 209–223. https://doi.org/10.2980/16-2-3220
10.2980/16-2-3220
Web of Science®Google Scholar
Browne, C. L., Paszkowski, C. A., Foote, A. L., Moenting, A., & Boss, S. M. (2009b). The relationship of amphibian abundance to habitat features across spatial scales in the Boreal Plains. Écoscience, 16(2), 209–223. https://doi.org/10.2980/16-2-3220
10.2980/16-2-3220
Web of Science®Google Scholar
Burnham, K. P., & Anderson, D. R. (2002). Model selection and multi-model inference: A practical information-theoretic approach. New York, NY: Springer-Verlag. https://doi.org/10.1007/b97636
Web of Science®Google Scholar
Calderon, M. R. (2019). Influence of water quality and habitat conditions on amphibian community metrics in rivers affected by urban activity. Urban Ecosystems, 22(4), 743–755. https://doi.org/10.1007/s11252-019-00862-w.
10.1007/s11252?019?00862?w
Web of Science®Google Scholar
Churko, G. (2016). Evaluating the landscape connectivity of five amphibian species using circuit theory (MSc Thesis). Eidgenössische Technische Hochschule Zürich. https://kbnl.ch/wp-content/uploads/2016/05/3.3_MasterThesis_Greg-Churko.pdf
Google Scholar
City of Calgary. (1994). Natural area management plan. Calgary Parks and Recreation Department. https://www.calgary.ca/content/dam/www/csps/parks/documents/planning-and-operations/natural-areas-and-wetlands/natural-area-management-plan.pdf
Google Scholar
City of Calgary. (2004). Calgary wetland conservation plan. City of Calgary Parks. https://www.calgary.ca/content/dam/www/csps/parks/documents/planning-and-operations/natural-areas-and-wetlands/wetland-conservation-plan.pdf
Google Scholar
City of Calgary. (2014). The city of Calgary biodiversity report. City of Calgary. https://www.calgary.ca/content/dam/www/csps/parks/documents/planning-and-operations/biodviersity-report-2014.pdf
Google Scholar
City of Calgary. (2015). Our BiodiverCity. Calgary's 10-year biodiversity strategic plan. CPS2015-0260. .https://www.calgary.ca/content/dam/www/csps/parks/documents/planning-and-operations/biodivercity-strategic-plan.pdf
Google Scholar
City of Calgary. (2017). The riparian action program: A blueprint for resilience. Calgary, Alberta, Canada. https://www.calgary.ca/content/dam/www/uep/water/documents/water-documents/riparian-action-program-report.pdf
Google Scholar
City of Calgary. (2021) City of Calgary's open data portal. https://data.calgary.ca/
Google Scholar
Clevenot, L., Carré, C., & Pech, P. (2018). A review of the factors that determine whether stormwater ponds are ecological traps and/or high-quality breeding sites for amphibians. Frontiers in Ecology and Evolution, 6, 1–12. https://doi.org/10.3389/fevo.2018.00040
10.3389/fevo.2018.00040
Web of Science®Google Scholar
Cook, C. N., Inayatullah, S., Burgman, M. A., Sutherland, W. J., & Wintle, B. A. (2014). Strategic foresight: How planning for the unpredictable can improve environmental decision-making. Trends in Ecology and Evolution, 29(9), 531–541. https://doi.org/10.1016/j.tree.2014.07.005
10.1016/j.tree.2014.07.005
PubMedWeb of Science®Google Scholar
Cooper, C. B., Hawn, C. L., Larson, L. R., Parrish, J. K., Bowser, G., Cavalier, D., Dunn, R. R., Haklay, M., Gupta, K. K., Jelks, N. O., Johnson, V. A., Katti, M., Leggett, Z., Wilson, O. R., & Wilson, S. (2021). Inclusion in citizen science: The conundrum of rebranding. Science, 372(6549), 1386–1388. https://doi.org/10.1126/science.abi6487
10.1126/science.abi6487
CASWeb of Science®Google Scholar
Cushman, S. A. (2006). Effects of habitat loss and fragmentation on amphibians: A review and prospectus. Biological Conservation, 128(2), 231–240. https://doi.org/10.1016/j.biocon.2005.09.031
10.1016/j.biocon.2005.09.031
Web of Science®Google Scholar
Cushman, S. A., & Landguth, E. L. (2012). Multi-taxa population connectivity in the northern Rocky Mountains. Ecological Modelling, 231, 101–112. https://doi.org/10.1016/j.ecolmodel.2012.02.011
10.1016/j.ecolmodel.2012.02.011
Web of Science®Google Scholar
Dodd, C. K. (2013). Frogs of the United States and Canada ( 2nd edn.). Johns Hopkins University Press.
10.1353/book.25108
Google Scholar
Garcia-Gonzalez, C., & Garcia-Vazquez, E. (2012). Urban ponds, neglected Noah's Ark for amphibians. Journal of Herpetology, 46(4), 507–514. https://doi.org/10.1670/10-227
10.1670/10?227
Web of Science®Google Scholar
Government of Alberta. (2013). Sensitive species inventory guidelines. Environment and sustainable resource development. Wildlife Management Department. https://open.alberta.ca/publications/sensitive-species-inventory-guidelines
Google Scholar
Groves, C. R., Jensen, D. B., Valutis, L. L., Redford, K. H., Shaffer, M. L., Scott, J. M., Baumgartner, J. V., Higgins, J. V., Beck, M. W., & Anderson, M. G. (2002). Planning for biodiversity conservation: Putting conservation science into practice. Bioscience, 52(6), 499–512. https://doi.org/10.1641/0006-3568(2002)052[0499:PFBCPC]2.0.CO;2
10.1641/0006-3568(2002)052[0499:PFBCPC]2.0.CO;2
Web of Science®Google Scholar
Hamer, A. J., & McDonnell, M. J. (2008). Amphibian ecology and conservation in the urbanising world: A review. Biological Conservation, 141(10), 2432–2449. https://doi.org/10.1016/j.biocon.2008.07.020
10.1016/j.biocon.2008.07.020
Web of Science®Google Scholar
Hamer, A. J., & Parris, K. M. (2011). Local and landscape determinants of amphibian communities in urban ponds. Ecological Applications, 21(2), 378–390. https://doi.org/10.1890/10-0390.1
10.1890/10-0390.1
PubMedWeb of Science®Google Scholar
Hamer, A. J., Smith, P. J., & McDonnell, M. J. (2012). The importance of habitat design and aquatic connectivity in amphibian use of urban stormwater retention ponds. Urban Ecosystems, 15(2), 451–471. https://doi.org/10.1007/s11252-011-0212-5
10.1007/s11252?011?0212?5
Web of Science®Google Scholar
Hammerson, G. (2008). Boreal chorus frog (Pseudacris maculata). NatureServe Explorer. https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.806592/Pseudacris_maculata
Google Scholar
Helldin, J. O., & Petrovan, S. O. (2019). Effectiveness of small road tunnels and fences in reducing amphibian roadkill and barrier effects at retrofitted roads in Sweden. PeerJ, 7, e7518. https://doi.org/10.7717/peerj.7518
10.7717/peerj.7518
PubMedWeb of Science®Google Scholar
Holtmann, L., Philipp, K., Becke, C., & Fartmann, T. (2017). Effects of habitat and landscape quality on amphibian assemblages of urban stormwater ponds. Urban Ecosystems, 20(6), 1249–1259. https://doi.org/10.1007/s11252-017-0677-y
10.1007/s11252?017?0677?y
Web of Science®Google Scholar
Hussain, Q. A. (2012). Global amphibian declines: A review. International Journal of Biodiversity and Conservation, 4(10), 348–357. https://academicjournals.org/journal/IJBC/article-abstract/683AC2810171
Google Scholar
Jarchow, C. J., Hossack, B. R., Sigafus, B. H., Schwalbe, C. R., & Muths, E. (2016). Modeling habitat connectivity to inform reintroductions: A case study with the Chiricahua leopard frog. Journal of Herpetology, 50(1), 63–69. https://doi.org/10.1670/14-172
10.1670/14?172
Web of Science®Google Scholar
Koen, E. L., Bowman, J., Sadowski, C., & Walpole, A. A. (2014). Landscape connectivity for wildlife: Development and validation of multispecies linkage maps. Methods in Ecology and Evolution, 5(7), 626–633. https://doi.org/10.1111/2041-210X.12197
10.1111/2041-210X.12197
Web of Science®Google Scholar
Konowalik, A., Najbar, A., Konowalik, K., Dylewski, Ł., Frydlewicz, M., Kisiel, P., Starzecka, A., Zaleśna, A., & Kolenda, K. (2020). Amphibians in an urban environment: A case study from a central European city (Wrocław, Poland). Urban Ecosystems, 23(2), 235–243. https://doi.org/10.1007/s11252-019-00912-3
10.1007/s11252?019?00912?3
Web of Science®Google Scholar
Lankau, H. (2015). Autonomous recording unit (ARU) deployment protocol. Version: 1 May 2015. http://bioacoustic.abmi.ca/resources/protocols
Google Scholar
Lee, T. S., Kahal, N. L., Kinas, H. L., Randall, L. A., Baker, T. M., Carney, V. A., Kendell, K., Sanderson, K., & Duke, D. (2021). Advancing amphibian conservation through citizen science in urban municipalities. Diversity, 13(211), 1–15. https://doi.org/10.3390/d13050211
10.3390/d13050211
Web of Science®Google Scholar
Linhoff, L. J., Soorae, P. S., Harding, G., Donnelly, M. A., Germano, J. M., Hunter, D. A., McFadden, M., Mendelson, L., Pessier, J. R., Sredl, M. J., & Eckstut, M. E. (2021). Guidelines for reintroductions and other conservation translocations. The Reintroduction and Invasive Species Specialist Groups’ Task Force on moving plants and animals for conservation purposes. Version 1.0. IUCN.https://portals.iucn.org/library/efiles/documents/2013-009.pdf
Google Scholar
MacKenzie, D. I., Nichols, J. D., Royle, J. A., Pollock, K. H., Bailey, L., & Hines, J. E. (2017). Occupancy estimation and modeling: Inferring patterns and dynamics of species occurrence ( 2nd edn.). Elsevier.
Google Scholar
Margules, C. R., & Pressey, R. L. (2000). Systematic conservation planning. Nature, 405(6783), 243–253. https://doi.org/10.1038/35012251
10.1038/35012251
CASPubMedWeb of Science®Google Scholar
Marsh, D. M., & Cosentino, B. J. (2019). Causes and consequences of non-random drop-outs for citizen science projects: Lessons from the North American amphibian monitoring program. Freshwater Science, 38(2), 292–302. https://doi.org/10.1086/701672
10.1086/701672
Web of Science®Google Scholar
McDonnell, M. J. (2015). Linking and promoting research and practice in the evolving discipline of urban ecology. Journal of Urban Ecology, 1(1), 1–6. https://doi.org/10.1093/jue/juv003
10.1093/jue/juv003
Google Scholar
McKinley, D. C., Miller-Rushing, A. J., Ballard, H. L., Bonney, R., Brown, H., Cook-Patton, S. C., Evans, D. M., French, R. A., Parrish, J. K., Phillips, T. B., Ryan, S. F., Shanley, L. A., Shirk, J. L., Stepenuck, K. F., Weltzin, J. F., Wiggins, A., Boyle, O. D., Briggs, R. D., Chapin, S. F., …, & Soukup, M. A. (2017). Citizen science can improve conservation science, natural resource management, and environmental protection. Biological Conservation, 208, 15–28. https://doi.org/10.1016/j.biocon.2016.05.015
10.1016/j.biocon.2016.05.015
Web of Science®Google Scholar
McKinley, D. C., Miller-rushing, A. J., Ballard, H. L., Bonney, R., Brown, H., Evans, D. M., French, R. A., Parrish, J. K., Phillips, T. B., Ryan, S. F., Shanley, L. A., Shirk, J. L., Stepenuck, K. F., Weltzin, J. F., Wiggins, A., Boyle, O. D., Briggs, R. D., Chapin III, S. F. C., Hewitt, D. A., …, & Soukup, M. A. (2015). Investing in citizen science can improve natural resource management and environmental protection. Issues in Ecology, 19, 1–27. https://pubs.er.usgs.gov/publication/70159470
PubMedWeb of Science®Google Scholar
McKinney, M. L. (2006). Urbanization as a major cause of biotic homogenization. Biological Conservation, 127(3), 247–260. https://doi.org/10.1016/j.biocon.2005.09.005
10.1016/j.biocon.2005.09.005
Web of Science®Google Scholar
McRae, B. H., Dickson, B. G., Timothy, H. K., & Shah, V. B. (2008). Using circuit theory to model connectivity in ecology, evolution, and conservation. Ecology, 89(10), 2712–2724. https://doi.org/10.1890/07-1861.1
10.1890/07-1861.1
PubMedWeb of Science®Google Scholar
Milanovich, J. R., Peterman, W. E., Barrett, K., & Hopton, M. E. (2012). Do species distribution models predict species richness in urban and natural green spaces? A case study using amphibians. Landscape and Urban Planning, 107(4), 409–418. https://doi.org/10.1016/j.landurbplan.2012.07
10.1016/j.landurbplan.2012.07
Web of Science®Google Scholar
Mushet, D. M., Euliss, N. H., & Stockwell, C. A. (2012). Mapping anuran habitat suitability to estimate effects of grassland and wetland conservation programs. Copeia, 2012(2), 321–330. https://doi.org/10.1643/ch-11-119
10.1643/ch?11?119
Web of Science®Google Scholar
Muths, E., Jung, R. E., Bailey, L. L., Adams, M. J., Corn, P. S., Dodd Jr. C. K., Fellers, G. M., Sadinski, W. J., Schwalbe, C. R., Walls, S. C., Fisher, R. N., Gallant, A. L., Battaglin, W. A., & Green, D. E. (2005). Amphibian Research and Monitoring Initiative (ARMI): A successful start to a national program in the United States. Applied Herpetology, 2, 355–371. https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1128&context=usgsstaffpub
10.1163/157075405774483139
Google Scholar
Nowakowski, A. J., Veiman-Echeverria, M., Kurz, D. J., & Donnelly, M. A. (2015). Evaluating connectivity for tropical amphibians using empirically derived resistance surfaces. Ecological Applications, 25(4), 928–942. https://doi.org/10.1890/14-0833.1
10.1890/14‐0833.1
PubMedWeb of Science®Google Scholar
Parris, K. M. (2006). Urban amphibian assemblages as metacommunities. Journal of Animal Ecology, 75(3), 757–764. https://doi.org/10.1111/j.1365-2656.2006.01096.x
10.1111/j.1365-2656.2006.01096.x
PubMedWeb of Science®Google Scholar
Pierce, J. R., Barton, M. A., Tan, M. M. J., Oertel, G., Halder, M. D., Lopez-Guijosa, P. A., & Nuttall, R. (2020). Actions, indicators, and outputs in urban biodiversity plans: A multinational analysis of city practice. PLOS One, 15(7), e0235773. https://doi.org/10.1371/journal.pone.0235773
10.1371/journal.pone.0235773
CASPubMedWeb of Science®Google Scholar
Randall, L. A., Smith, D. H. V., Jones, B. L., Prescott, D. R. C., & Moehrenschlager, A. (2015). Seasonal differences in extinction and colonization drive occupancy dynamics of an imperilled amphibian. PloS One, 10(5), e0127059. https://doi.org/10.1371/journal.pone.0127059
10.1371/journal.pone.0127059
PubMedWeb of Science®Google Scholar
Randall, L., Lloyd, N., & Moehrenschlager, A. (2018). Guidelines for mitigation translocations of amphibians: Applications for Canada's prairie provinces. Version 1.0. Center for Conservation Research. Calgary Zoological Society. https://www.calgaryzoo.com/sites/default/files/2018-06/amphibian_translocation_guidelines.pdf
Google Scholar
Rubbo, M. J., & Kiesecker, J. M. (2005). Amphibian breeding distribution in an urbanized landscape. Conservation Biology, 19(2), 504–511. https://doi.org/10.1111/j.1523-1739.2005.000101.x
10.1111/j.1523-1739.2005.000101.x
Web of Science®Google Scholar
Sanderson, K. (2022a). A framework to identify priority wetland habitats and movement corridors for urban amphibian conservation - Raw Data. Zenodo. https://doi.org/10.5281/zenodo.6251519
Google Scholar
Sanderson, K. (2022b). A framework to identify priority wetland habitats and movement corridors for urban amphibian conservation – Amphibian Modelling Datasets. Zenodo. https://doi.org/10.5281/zenodo.6233027
Google Scholar
Sandström, U. G., Angelstam, P., & Khakee, A. (2006). Urban comprehensive planning — identifying barriers for the maintenance of functional habitat networks. Landscape and Urban Planning, 75(1–2), 43–57. https://doi.org/10.1016/j.landurbplan.2004.11.016
10.1016/j.landurbplan.2004.11.016
Web of Science®Google Scholar
Sarkar, S., Pressey, R. L., Faith, D. P., Margules, C. R., Fuller, T., Stoms, D. M., Moffett, A., Wilson, K. A., Williams, K. J., Williams, P. H., & Andelman, S. (2006). Biodiversity conservation planning tools: Present status and challenges for the future. Annual Review of Environment and Resources, 31(1), 123–159. https://doi.org/10.1146/annurev.energy.31.042606.085844
10.1146/annurev.energy.31.042606.085844
Web of Science®Google Scholar
Scheffers, B. R., & Paszkowski, C. A. (2012). The effects of urbanization on North American amphibian species: Identifying new directions for urban conservation. Urban Ecosystems, 15(1), 133–147. https://doi.org/10.1007/s11252-011-0199-y
10.1007/s11252?011?0199?y
Web of Science®Google Scholar
Scheffers, B. R., & Paszkowski, C. A. (2013). Amphibian use of urban stormwater wetlands: The role of natural habitat features. Landscape and Urban Planning, 113, 139–149. https://doi.org/10.1016/j.landurbplan.2013.01.001
10.1016/j.landurbplan.2013.01.001
Web of Science®Google Scholar
Scherer, R. D., Muths, E., & Noon, B. R. (2012). The importance of local and landscape-scale processes to the occupancy of wetlands by pond-breeding amphibians. Population Ecology, 54(4), 487–498. https://doi.org/10.1007/s10144-012-0324-7
10.1007/s10144?012?0324?7
Web of Science®Google Scholar
Schwartz, M. W. (2006). How conservation scientists can help develop social capital for biodiversity. Conservation Biology, 20(5), 1550–1552. https://doi.org/10.1111/j.1523-1739.2006.00421.x
10.1111/j.1523?1739.2006.00421.x
PubMedWeb of Science®Google Scholar
Schwartz, M. W., Cook, C. N., Pressey, R. L., Pullin, A. S., Runge, M. C., Salafsky, N., Sutherland, W. J., & Williamson, M. A. (2018). Decision support frameworks and tools for conservation. Conservation Letters, 11(2), 1–12. https://doi.org/10.1111/conl.12385
10.1111/conl.12385
Web of Science®Google Scholar
Shih, W. Y., Mabon, L., & Puppim de Oliveira, J. A. (2020). Assessing governance challenges of local biodiversity and ecosystem services: Barriers identified by the expert community. Land Use Policy, 91, 104291. https://doi.org/10.1016/j.landusepol.2019.104291
10.1016/j.landusepol.2019.104291
Web of Science®Google Scholar
Shirk, J. L., Ballard, H. L., Wilderman, C. C., Phillips, T., Wiggins, A., & Jordan, R. (2012). Public participation in scientific research: A framework for deliberate design. Ecology and Society, 17(2), 187–201. https://doi.org/10.5751/ES-04705-170229
10.5751/ES?04705?170229
Web of Science®Google Scholar
Shulse, C. D., Semlitsch, R. D., Trauth, K. M., & Williams, A. D. (2010). Influences of design and landscape placement parameters on amphibian abundance in constructed wetlands. Wetlands, 30(5), 915–928. https://doi.org/10.1007/s13157-010-0069-z
10.1007/s13157-010-0069-z
Web of Science®Google Scholar
Shwartz, A., Turbé, A., Julliard, R., Simon, L., & Prévot, A. C. (2014). Outstanding challenges for urban conservation research and action. Global Environmental Change, 28(1), 39–49. https://doi.org/10.1016/j.gloenvcha.2014.06.002
10.1016/j.gloenvcha.2014.06.002
Web of Science®Google Scholar
Simon, J. A., Snodgrass, J. W., Casey, R. E., & Sparling, D. W. (2009). Spatial correlates of amphibian use of constructed wetlands in an urban landscape. Landscape Ecology, 24(3), 361–373. https://doi.org/10.1007/s10980-008-9311-y
10.1007/s10980?008?9311?y
Web of Science®Google Scholar
Sinsch, U. (1990). Migration and orientation in anuran amphibians. Ethology, Ecology and Evolution, 2, 65–79. https://doi.org/10.1080/08927014.1990.9525494
10.1080/08927014.1990.9525494
Web of Science®Google Scholar
Skidds, D. E., Golet, F. C., Paton, P. W. C., & Mitchell, J. C. (2007). Habitat correlates of reproductive effort in Wood Frogs and Spotted Salamanders in an urbanizing watershed. Journal of Herpetology, 41(3), 439–450. https://doi.org/10.1670/0022-1511(2007)41[439:hcorei]2.0.co;2
10.1670/0022?1511(2007)41[439:hcorei]2.0.co;2
Web of Science®Google Scholar
Smith, M. A., & Green, D. M. (2005). Dispersal and the metapopulation paradigm in amphibian ecology and conservation: Are all amphibian populations metapopulations? Ecography, 28(1), 110–128. https://doi.org/10.1111/j.0906-7590.2005.04042.x
10.1111/j.0906-7590.2005.04042.x
Web of Science®Google Scholar
Soorae, P. S. (Ed.). (2016). Global re-introduction perspectives: 2016. Case-studies from around the globe. IUCN/SSC Re- introduction Specialist Group and Abu Dhabi, AE. Environment Agency.
Google Scholar
Sterling, E. J., Betley, E., Sigouin, A., Gomez, A., Toomey, A., Cullman, G., Malone, C., Pekor, A., Arengo, F., Blair, M., Filardi, C., Landrigan, K., & Porzecanski, A. L. (2017). Assessing the evidence for stakeholder engagement in biodiversity conservation. Biological Conservation, 209, 159–171. https://doi.org/10.1016/j.biocon.2017.02.008
10.1016/j.biocon.2017.02.008
Web of Science®Google Scholar
Sterrett, S. C., Katz, R. A., Brand, A. B., Fields, W. R., Dietrich, A. E., Hocking, D. J., Foreman, T. M., Wiewel, A. N. M., & Campbell Grant, E. H. (2019). Proactive management of amphibians: Challenges and opportunities. Biological Conservation, 236, 404–410. https://doi.org/10.1016/j.biocon.2019.05.057
10.1016/j.biocon.2019.05.057
Web of Science®Google Scholar
Turo, K. J., & Gardiner, M. M. (2020). The balancing act of urban conservation. Nature Communications, 11(1), 1–5. https://doi.org/10.1038/s41467-020-17539-0
10.1038/s41467‐020‐17539‐0
PubMedWeb of Science®Google Scholar
Wade, A. A., McKelvey, K. S., & Schwartz, M. K. (2015). Resistance-surface-based wildlife conservation connectivity modeling: Summary of efforts in the united states and guide for practitioners. USDA Forest Service - General Technical Report RMRS-GTR, 2015(333), 1–93. https://doi.org/10.2737/RMRS-GTR-333
10.2737/RMRS?GTR?333
Google Scholar
Warton, D. I., & Hui, F. K. C. (2011). The arcsine is asinine: The analysis of proportions in ecology. Ecology, 92(1), 3–10. https://doi.org/10.1890/10-0340.1
10.1890/10-0340.1
PubMedWeb of Science®Google Scholar
Westgate, M. J., Scheele, B. C., Ikin, K., Hoefer, A. M., Beaty, R. M., Evans, M., Osborne, W., Hunter, D., Rayner, L., & Driscoll, D. A. (2015). Citizen science program shows urban areas have lower occurrence of frog species, but not accelerated declines. Plos One, 10(11), e0140973. https://doi.org/10.1371/journal.pone.0140973
10.1371/journal.pone.0140973
PubMedWeb of Science®Google Scholar
Zaffaroni, M., Zamberletti, P., Creed, I. F., Accatino, F., De Michele, C., & DeVries, B. (2019). Safeguarding wetlands and their connections within wet landscapes to improve conservation outcomes for threatened amphibian species. Journal of the American Water Resources Association, 55, 641–656. https://doi.org/10.1111/1752-1688.12751
10.1111/1752?1688.12751
Web of Science®Google Scholar
Zamberletti, P., Zaffaroni, M., Accatino, F., Creed, I. F., & De Michele, C. (2018). Connectivity among wetlands matters for vulnerable amphibian populations in wet landscapes. Ecological Modelling, 384, 119–127. https://doi.org/10.1016/j.ecolmodel.2018.05.008
10.1016/j.ecolmodel.2018.05.008
Web of Science®Google Scholar
Zari, M. P. (2018). The importance of urban biodiversity — an ecosystem services approach. Biodiversity International Journal, 2(4), 357–360. https://doi.org/10.15406/bij.2018.02.00087
10.15406/bij.2018.02.00087
Google Scholar

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Volume3, Issue2

April–June 2022

e12139

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Key Themes in Ecology – A Cross-Journal Virtual Special Issue in Support of the British Ecological Society 2022 Annual Meeting

A framework to identify priority wetland habitats and movement corridors for urban amphibian conservation

Abstract

1 INTRODUCTION

2 MATERIALS AND METHODS