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International Journal of Ecology
Volume 2012 (2012), Article ID 989872, 11 pages
A Place to Call Home: Amphibian Use of Created and Restored Wetlands
1Department of Biology, Texas State University-San Marcos, 601 University Drive, San Marcos, TX 78666, USA
2Department of Integrative Biology, University of Guelph, 50 Stone Road East, Guelph, ON, Canada N1G 2W1
3Center for Restoration of Ecosystems and Watersheds, School of Civil Engineering and Environmental Science, University of Oklahoma, 202 West Boyd Street, Norman, OK 73019, USA
Received 8 July 2011; Revised 26 September 2011; Accepted 20 October 2011
Academic Editor: L. Smith
Copyright © 2012 Donald J. Brown et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Loss and degradation of wetland habitats are major contributing factors to the global decline of amphibians. Creation and restoration of wetlands could be a valuable tool for increasing local amphibian species richness and abundance. We synthesized the peer-reviewed literature addressing amphibian use of created and restored wetlands, focusing on aquatic habitat, upland habitat, and wetland connectivity and configuration. Amphibian species richness or abundance at created and restored wetlands was either similar to or greater than reference wetlands in 89% of studies. Use of created and restored wetlands by individual species was driven by aquatic and terrestrial habitat preferences, as well as ability to disperse from source wetlands. We conclude that creating and restoring wetlands can be valuable tools for amphibian conservation. However, the ecological needs and preferences of target species must be considered to maximize the potential for successful colonization and long-term persistence.
1. Amphibian Habitat Loss and Restoration
1.1. Amphibian Decline
It is widely accepted that amphibians are experiencing world-wide declines in abundance and diversity (e.g., [1–5]). Stuart et al.  estimated more than 2400 of the approximately 5700 species had experienced severe population declines or extinction, and there is little evidence suggesting these trends have improved in recent years . Primary reasons for declines are summarized by Collins and Storfer  and include the spread of invasive species, increasing infectious disease outbreaks, patterns of global climate change, and human land use practices. Of these, human land use is arguably one of the most readily identifiable negative impacts on amphibian populations.
Wetland habitats are often drained and altered to accommodate development of agriculture and urban expansion, with devastating effects on local amphibian populations [8–10]. According to Mitsch and Gosselink , the decline in the number of wetlands across the world varies by region from 33% to more than 90%. Habitat loss and fragmentation may exacerbate the negative impacts associated with other causes of declines such as habitat degradation, resulting in decreased mating success and increased susceptibility to other biotic and abiotic factors (e.g., [7, 9, 12–14]).
1.2. Creation and Restoration of Wetland Habitat
In recognition of the importance of wetland ecosystems to both biotic (e.g., species richness, food chain and biodiversity support, habitat provision) and abiotic (e.g., elemental cycling, hydrologic buffering, climate stabilization) processes, numerous policies and regulations have been enacted to mitigate the loss of wetland habitat. The United States has actively pursued the restoration of wetland habitat through a de facto national policy of “no net loss,” although no national wetland protection law exists and these ecosystems are regulated by a suite of land use and water quality regulations. Wetland protection policies have evolved in the past two decades from simple area replacement strategies to incorporation of methods that attempt to evaluate lost and subsequently restored or created function. Principal among these attempts to move beyond simple area replacement is the Hydrogeomorphic approach to assessing wetlands functions (HGM; [15–17]). The HGM approach regionalizes wetland functional assessment and incorporates reference wetland condition assessments based on geomorphic position and hydrologic characteristics.
Estimating the ecological success of created and restored wetlands is complex, multifaceted, and site- and project-specific. Compared to analyses of area replacement, useful analyses of ecological functional replacement require considerable time, financial resources, and ecological expertise [18–20]. Little information exists concerning whether the ecosystem functions of created and restored sites sufficiently compensate for those lost in the original wetlands . Moreover, the ecological goals of wetland creation and restoration projects are often unclear or inappropriate for a given restoration project design, producing conflicting results and unsuccessful restoration attempts [22, 23].
Efforts to identify appropriate reference wetlands for comparative evaluation may not follow established scientific protocols or be simply overlooked or never undertaken [22, 24]. In the United States, some attempts have been made to combine the HGM approach with abundance estimates of aquatic bioindicator species using various indices of biotic integrity (IBIs; [25–27]). Given the controversy surrounding assessment approaches, and thus determination of reference conditions, Stein et al.  argue that wetland management needs should drive the selected approach, and not the other way around. Finally, monitoring of created and restored wetlands is frequently inadequate, and when conducted primarily focuses on wetland hydrology, biogeochemistry, and vegetation, with little emphasis on faunal use and abundance . Even when faunal colonization is an explicit restoration objective, assessment procedures often fail to recognize the importance of nontarget species richness and related ecological function [30, 31]. Given the importance of faunal activity to healthy wetland function , this dearth of information suggests a need for evaluative studies of faunal use of created and restored wetlands.
2. Important Considerations for Amphibians
2.1. Aquatic Habitat
For most amphibians with complex life cycles , standing or slow-moving water is necessary for the egg and tadpole development stages . Thus, aquatic habitat quality can be an important determinant of amphibian species composition, richness, and abundance [34, 35]. Wetland-breeding amphibian species vary substantially in developmental timing strategies and aquatic habitat preferences . However, several aquatic habitat features generally appear to benefit amphibians, including lack of predatory fish [36, 37], lack of eutrophication [38, 39], and presence of aquatic macrophytes [40, 41]. Macrophytes increase habitat complexity, and thus can reduce predation pressure by creating refuge zones for larval amphibians [42, 43]. Although nutrients are important for phytoplankton growth , which is a primary food source used during the tadpole stage for most wetland-breeding species [33, 45], many amphibians are sensitive to high levels of ammonium and nitrate, low pH, and low levels of dissolved oxygen [46–48]. Further, eutrophication can potentially increase pathogenic infections .
Wetland hydroperiod is also important . Long periods of standing water could potentially result in high species richness because species with both short and long developmental periods could utilize the wetland. However, many species have evolved to breed in ephemeral water bodies, and some typically will not utilize wetlands with long hydroperiods , which are vulnerable to fish colonization and increased interspecific competition and predation [51–53]. For example, a positive correlation between amphibian species richness and depressional wetland hydroperiod was observed in the southeastern United States, but many species adapted to ephemeral wetlands were not found in wetlands with long hydroperiods .
2.2. Upland Habitat
Wetland-breeding amphibians require suitable aquatic and terrestrial habitat for long-term persistence . Based on 21 species investigations, anuran (frog and toad) home range sizes were between 1 m2 and 1,900 m2, with a median of 40 m2 . Based on 13 species investigations, salamander home range sizes were between 0.1 m2 and 90 m2, with a median of 4 m2 . Rittenhouse and Semlitsch  found that 50% and 95% of amphibian species investigated () remained within 93 m and 664 m, respectively, of wetlands during nonbreeding seasons. Many amphibians are specialized for particular upland habitat types, and populations will not persist in suboptimal habitat (e.g., 17 amphibians are endemic to longleaf pine (Pinus palustris) savanna ecosystems in the southeastern United States; ). Thus, upland habitat can be critically important to long-term persistence of amphibian populations.
2.3. Wetland Connectivity and Configuration
Population sizes naturally fluctuate for many wetland-breeding amphibian species, primarily in relation to annual weather variability [2, 57]. Colonization from surrounding water bodies is often important for long-term persistence of populations and subpopulations [58, 59]. Further, because probability of colonization is inversely related to distance traveled , the establishment of several wetlands in close proximity to one another is typically optimal for long-term persistence . However, amphibians are vulnerable to road mortality, and thus wetland complexes that are bisected by roads can be problematic [14, 61, 62].
While in general increasing wetland density and connectivity benefits amphibians, there are situations where this can run counter to conservation goals. Florance et al.  concluded that artificial water points (e.g., cattle troughs) served as dry season refuges for invasive cane toads (Bufo [Rhinella] marinus) in Australia, which aided in range expansion of this species. Gaston et al.  showed that probability of reproduction in the endangered Houston toad (Bufo [Anaxyrus] houstonensis) increased exponentially with number of calling males, and indicated that increasing wetland density in suboptimal habitat could negatively impact this species by decreasing toad density at individual wetlands. Thus, careful consideration of the placement of wetlands within the surrounding landscape is warranted.
3. Amphibian Use of Created and Restored Wetlands
For the purpose of this review, we included only peer-reviewed studies. We recognized created and restored wetlands as distinct from enhanced and treatment wetlands built specifically for water quality improvement, per Mitsch and Jørgensen , and focused only on those studies that addressed the conversion of an existing upland or shallow-water area to wetland habitat (created), or an attempt to return a wetland to a previously occurring wetland condition (restored). We omitted studies that addressed improvement of existing wetland function (enhancement) or creation of new wetlands for water quality improvement and contaminant removal (constructed/treatment). This distinction permitted us to focus on studies addressing the mitigation of wetland habitat loss due to development or environmental change.
It bears noting that treatment wetlands have been shown to support diverse populations of amphibians and may be viable replacement habitat (e.g., [65, 66]). However, treatment wetlands for polluted waters do not typically consider faunal use in their designs, and may actively restrict wildlife from the site using exclusion barriers, trapping, and other habitat modifications that promote water quality improvement while reducing the presence of wildlife . Moreover, this topic has received little attention in the literature. As such, the use of treatment wetlands as amphibian habitat necessarily fell outside the scope of this review.
We found 37 peer-reviewed articles that explicitly addressed amphibian use of created or restored wetlands (Table 1). Twenty-six of the studies included controls in their investigation, which were either reference wetlands or historic survey data. The majority of studies were conducted in the United States () and surrounding land cover types were primarily forest or agriculture. Species richness or abundance of some or all species was greater at created or restored wetlands versus reference wetlands in 54% of studies, similar in 35% of studies, and lower in 11% of studies.
3.1. Aquatic Habitat
Thirty-three studies addressed the influence of aquatic habitat differences on wetland creation or restoration success. Two aquatic habitat variables were discussed in nearly all of the papers, presence or absence of fish and wetland hydroperiod. With the exception of anurans in the families Ranidae and Bufonidae, amphibian occupancy and abundance was typically negatively associated with presence of fish [41, 60, 68–74], especially for newts [75–77]. In particular, sunfish (Lepomis spp. [41, 60, 69, 71, 73, 75]), goldfish (Carassius auratus [41, 76, 77]), and largemouth bass (Micropterus salmoides [41, 60, 73]) appeared to negatively influence amphibians. The apparent positive association between Rana spp. and fish presence could be primarily consequent of both taxa utilizing permanent wetlands [29, 78, 79]. However, studies have shown that fish can increase tadpole survivorship of Rana and Bufo through predatory invertebrate reduction [80, 81], as well as through reduction of predatory amphibian larvae . In addition, amphibian larval palatability influences predation levels, and both Bufo spp. and Rana spp. have been found to be unpalatable to some fish species [83, 84]. Finally, gape-limitations of smaller-bodied fish can reduce predation on amphibian larvae .
Created and restored wetlands were typically larger, deeper, and had longer hydroperiods than natural wetlands, and in general larger wetlands with longer hydroperiods resulted in greater species richness [86–90]. Based on seven survey years at 10 constructed and 10 reference wetlands, Petranka et al.  found that occupancy did not differ between created and reference wetlands for the four salamander species investigated. Further, although occupancy at created wetlands was lower for the eastern newt (Notophthalmus viridescens), it was significantly higher for the six anurans investigated. Brand and Snodgrass  investigated use of created and natural wetlands in suburban and forested landscapes by six anurans over two survey years. Calling activity occurred almost exclusively in created wetlands, and larvae were found only in created wetlands. Brand and Snodgrass  speculated that the short hydroperiods in natural wetlands in their system would prevent reproductive success, possibly due to changes in natural hydrology caused by surrounding landscape alterations.
Although most of the studies found the longer hydroperiods associated with created wetlands to be positively associated with amphibian use, the observed relation was sometimes dependent on the wetlands being free of fish. For example, Julian et al.  found that spotted salamander (Ambystoma maculatum) and wood frog (Rana [Lithobates] sylvatica) egg masses were less likely to be found in created than natural wetlands, and suggested the primary driver of this result was presence of fish in created wetlands. In most cases wetlands with an intermediate hydroperiod (i.e., wetlands that hold water for several months each year but are not permanent) had the highest species richness because they allowed most amphibian taxa to complete larval development while minimizing fish colonization [88, 91].
In most studies, amphibian species richness and abundance was positively associated with presence and abundance of emergent vegetation [41, 68, 74, 77, 89, 92, 93]. Presence of emergent vegetation was positively associated with reproductive success of the Columbia spotted frog (Rana luteiventris) and eastern long-toed salamander (Ambystoma macrodactylum) at created ponds in Idaho . Crested newts (Triturus cristatus) were more likely to colonize wetlands containing submerged aquatic vegetation . Lesbarrères et al.  determined that number of vegetation strata positively influenced species richness and diversity at created ponds in France. Green frog (Rana [Lithobates] clamitans) occupancy of restored wetlands in Canada was positively influenced by percent cattail (Typha spp. ). However, Porej and Hetherington  found that amount of emergent vegetation did not influence amphibian species richness at created wetlands in Ohio, USA. Similarly, Lehtinen and Galatowitsch  determined that aquatic vegetation cover did not influence amphibian species richness at restored wetlands in Minnesota, USA.
Several studies addressed the influence of slope on amphibian use of created and restored wetlands [71, 74, 77]. The presence of a shallow littoral zone was positively associated with amphibian species richness at created wetlands in Ohio, USA . Shulse et al.  determined that relative abundance of American toads (Bufo [Anaxyrus] americanus) and boreal chorus frogs (Pseudacris maculata) at created wetlands in Missouri, USA, was negatively associated with wetland slope. However, Rannap et al.  did not find a significant relationship between either wetland slope or width of the shallow littoral zone and relative abundance of crested newts or common spadefoot toads (Pelobates fuscus) in restored ponds.
3.2. Upland Habitat
Ten studies assessed the influence of surrounding upland habitat on wetland creation or restoration success, with only two failing to detect an influence of the upland habitats. Bowers et al.  found that tree planting in and near riparian zones did not influence amphibian species richness for species during initial restoration stages at the Savannah River Site in South Carolina, USA. Petranka et al.  determined that distance from forest cover did not influence colonization rates or species richness at a mitigation site in North Carolina, USA. Although the remaining studies determined that adjacent upland habitat was important, results were dependent upon habitat preferences of the species investigated. Brand and Snodgrass  found that wood frogs used forested wetlands more than suburban wetlands, but in general, species richness was greater at suburban wetlands. Crested newts preferred golf course ponds, whereas smooth newts (Triturus vulgaris) preferred adjacent parklands . Simon et al.  found that forest cover within 500 m of wetlands was a better predictor of amphibian species richness in Maryland, USA, than differences among created wetlands. Shulse et al.  found that salamander and spring peeper (Pseudacris crucifer) abundances were negatively associated with percent cropland in the surrounding landscape, and Monello and Wright  found that distance from agricultural land was positively associated with Columbia spotted frog (Rana luteiventris) reproduction.
Three studies showed that roads adjacent to created wetlands negatively influenced presence of amphibians [74, 87, 97]. Conversely, Petranka et al.  did not detect an association between species richness or number of egg masses and distance to paved roads for wood frogs and spotted salamanders at created ponds in North Carolina, USA, and Balcombe et al.  noted that proximity of created wetlands to major roads did not seem to negatively affect anuran abundance. Pechmann et al.  and Lehtinen and Galatowitsch  speculated that roads may have acted as dispersal barriers for colonization of created wetlands in South Carolina, USA, and restored wetlands in Minnesota, USA, respectively.
3.3. Wetland Connectivity and Configuration
Sixteen studies addressed the role of connectivity and configuration on wetland creation or restoration success. These studies assessed the influence of availability or density of source wetlands, but did not explicitly investigate the influence of spatial arrangement. Newt colonization was heavily dependent on source wetlands in close proximity [68, 75–77, 82]. In a study that investigated both local scale (e.g., hydroperiod, fish presence) and landscape scale (e.g., elevation, wetland density) variables, wetland connectivity was the most important variable for predicting high species richness . Lehtinen and Galatowitsch  determined that distance to source ponds was an important factor in predicting amphibian species richness and speculated that the lack of colonization for four species was due to poor dispersal abilities. Shulse et al.  found that surrounding pond density or percent wetland in the surrounding landscape was positively associated with abundances of five amphibian species at created wetlands in Missouri, USA. Wetland complexes with variable hydrologic regimes were found to increase potential for restoration success by catering to species-specific preferences and buffering effects of weather variability and disease outbreaks [60, 73, 77, 82, 98]. Further, Petranka and Holbrook  indicated that a “patchy population” wetland complex design, characterized by large variability in wetland size, hydroperiod, and spatial proximity, was better than a metapopulation design. A patchy population design allows for adaptive habitat switching, thus maintaining a high probability of population persistence within the wetland complex.
Habitat corridors can be important for restoring or maintaining wetland connectivity, and several studies considered the influence of corridors in their investigations. Rannap et al.  found that created wetlands surrounded by forest cover were colonized by amphibians from source wetlands more quickly than those surrounded by meadows. Vasconcelos and Calhoun  documented wood frog and spotted salamander movement patterns to and from restored wetlands in Maine, USA, and found that both species preferred to move through forested habitat compared to meadows. Lee et al.  determined that type of corridor influenced connectivity among restored wetlands in Taiwan, with wet areas containing dense vegetation being the most used, and drier meadows near roads being the least used. Chovanec et al.  investigated amphibian use of restored riparian areas along a heavily modified stretch of the Danube River in Austria. They found wetlands hydrologically isolated from the river had greater amphibian species richness and a larger number of successfully breeding species, but corridors along the river improved landscape-scale wetland connectivity. Bowers et al.  investigated amphibian colonization of a restored bottomland hardwood forest corridor in South Carolina, USA, and found no differences in relative abundance or diversity between restored and nonrestored corridors three years after the restoration efforts. However, the authors predicted that restoration attempts would be successful in the long term, as the restored plant community developed into a mature forest. Finally, several studies indicated that upland habitat composition was important for connectivity among wetlands [71, 75, 76, 100].
4. Species-Specific Responses to Wetland Creation and Restoration
In this section we summarize species-specific responses that have been observed for North American amphibians. To ensure broad applicability, we limited our review to species that were explicitly considered in at least eight papers.
4.1. American Toad
American toads (Bufo [Anaxyrus] americanus) are habitat generalists found throughout much of the eastern United States and Canada [101, 102]. Of the 15 studies we reviewed that investigated the American toad, only one did not find the species utilizing created or restored wetlands, in this case man-made bog pools . However, it also was not found at reference wetlands included in the study, likely due to acidic conditions . Brand and Snodgrass  found that American toads exclusively used stormwater and created wetlands, which was likely related to their larger sizes and reported longer hydroperiods than natural adjacent wetlands. However, Brand and Snodgrass  speculated that upland habitat alteration had altered the hydrology of the natural wetlands. This species colonized created and restored wetlands rapidly [104, 105], and relative abundances in most studies were similar to reference wetlands [29, 92]. Unlike many other amphibians, presence of predatory fish did not seem to negatively influence use of created and restored wetlands by American toads [71, 73, 74]. American toad tadpoles have been shown to be toxic to some fish and invertebrate predators [106–108], and thus wetlands containing fish may not be highly detrimental to reproductive success. However, Petranka and Holbrook  found that American toads avoided restored ponds containing wood frog tadpoles, likely due to their predatory nature . Finally, this species appeared to prefer ponds with shallow slopes [71, 74].
Bullfrogs (Rana [Lithobates] catesbeiana) have one of the largest natural ranges of North American amphibians, stretching from northern Mexico across the central and eastern United States to southern Canada . However, the species has also been introduced throughout much of the western United States, and is potentially contributing to declines of several amphibian species . Indeed, three of the studies reviewed here included bullfrogs outside of their natural range [41, 79, 91]. Of the 14 studies we reviewed that investigated the bullfrog, all of them reported bullfrog use of created and restored wetlands. Wetlands with long hydroperiods (i.e., wetlands that do not typically dry every year) are necessary for successful bullfrog reproduction due to a long larval stage , and created wetlands typically had longer hydroperiods than natural wetlands [71, 74, 86]. Bullfrog use of created and restored wetlands was positively associated with pond depth , and density of wetlands in the surrounding landscape . Presence of fish did not deter wetland use [41, 71, 91].
4.3. Green Frog
The distribution of green frogs (Rana [Lithobates] clamitans) extends throughout the eastern United States and southern Canada . This species selects wetlands with long hydroperiods due to a long larval stage, as well as adult habitat preference for aquatic environments . Of the 16 studies we reviewed that investigated the green frog, only one showed that it did not use created or restored wetlands, possibly due to limited dispersal ability . Indeed, several studies reported delayed wetland colonization for green frogs [50, 90]. Wetland use was positively associated with hydroperiod and density of surrounding wetlands, and presence of fish did not deter use [71, 73, 74]. Green frog relative abundance was typically higher at created and restored wetlands than reference wetlands, potentially due to longer hydroperiods in the created and restored wetlands [29, 86, 88, 93].
4.4. Wood Frog
Wood frogs (Rana [Lithobates] sylvatica) have the largest distribution of any North American amphibian, extending from the east-central United States to northern Canada and west to Alaska [101, 102]. This species is an explosive breeder with a short larval period, which allows it to breed in ephemerally flooded pools . Outside of the breeding season, the wood frog resides in the surrounding terrestrial environment, and strongly prefers forested habitat [112, 113]. Of the 13 studies we reviewed that investigated the wood frog, all of them reported wood frog use of created and restored wetlands. Wood frogs were found to rapidly colonize created and restored wetlands [60, 104]. In most studies this species used created and restored wetlands more than natural wetlands, which were typically larger and had longer hydroperiods [86–88, 93]. However, wood frogs showed a strong aversion to wetlands inhabited by fish [72, 78], and abandoned created wetlands after fish colonization .
4.5. Spotted Salamander
Spotted salamanders (Ambystoma maculatum) are found across the eastern United States and southeastern Canada . This species occupies terrestrial habitat when not engaged in breeding activity, and prefers forested environments [101, 113]. Of the 10 studies we reviewed that investigated the spotted salamander, only one did not find the species utilizing created or restored wetlands. However, detection at the reference site consisted of only one individual . Use of created, restored, and reference wetlands varied among the studies [71, 72, 86]. However, in all cases presence and relative abundance of the spotted salamander was negatively associated with the presence of fish [60, 71–74].
5. Recommendations for Wetland Creation and Restoration
Currently the literature on amphibian use of created and restored wetlands is limited to a small number of species, primarily in North America. However, based on these studies, wetland creation and restoration may be effective for enhancing amphibian abundance and diversity, and thus may be a valuable tool for mitigating amphibian population declines . There was no indication that “artificial” wetlands were inherently less suitable for amphibian use than natural wetlands. Rather, amphibian occupancy and abundance was strongly related to species-specific habitat associations and requirements, as well as dispersal ability.
These studies indicate that needs and preferences of target species should be a major consideration in wetland creation and restoration [77, 91, 116, 117]. Wetlands that are constructed or restored with the goal of providing high-quality habitat for amphibians must consider both the aquatic and surrounding terrestrial habitat, as well as colonization potential. The uplands surrounding managed wetlands are often referred to as “buffer zones,” and are typically ≤30 m in width surrounding the wetland for those areas wherein protective legislation exists . Buffer zones ≤30 m are clearly not sufficient for most anurans, which require 100 m or more [33, 55]. Based on empirical habitat use investigations, Semlitsch  suggested that buffer zones for salamander populations should extend at least 164 m from the edge of a wetland. Further, Semlitsch and Bodie  determined that core habitat zones for anurans were between 205 m and 368 m from the edge of a wetland. Thus, it is clear that maximizing the value of wetland creation to amphibians will require the integration of policy concerning surrounding upland habitat.
In addition to protecting habitat around wetlands from human development (e.g., buildings and roads), the habitat structure of surrounding uplands is important and should be managed for target species [100, 121, 122]. For most threatened and endangered amphibians, and indeed most wildlife species, habitat loss and degradation are principle drivers of long-term declines [3, 5, 123]. Barring disease outbreaks [124, 125], amphibian species that are of greatest conservation concern are typically habitat specialists unable to adapt to human-influenced terrestrial or aquatic habitat changes [126–128]. Thus, preserving or restoring upland systems can be essential for long-term success of wetland restoration programs, and the influence of upland habitat on wetland connectivity should be explicitly considered in restoration programs [93, 129].
It is apparent from this review that there is still much knowledge to gain concerning creation and restoration of wetlands for the benefit of amphibians. We found that many studies were observational in nature, and lacked rigorous experimental design or statistical frameworks. Although this was not surprising given the studies were conducted in real systems with corresponding experimental limitations, the variability in experimental design and data collection made it impossible for us to analyze these data using meta-analysis techniques. Despite these limitations, we believe the following patterns emerged from these studies, which are useful for assisting with future wetland creation and restoration efforts: (1) colonization was influenced by proximity to source wetlands (a function of dispersal capability) and upland habitat connectivity (a function of habitat selection); (2) wetlands with intermediate hydroperiods supported the greatest number of species; (3) presence of aquatic vegetation and shallow slopes increased amphibian use; (4) presence of fish decreased use for most amphibians; and (5) positive results from breeding habitat creation were apparent in the short-term (typically within one to two years), whereas upland and corridor habitat restoration projects required longer time periods to be effective, particularly in forested habitats.
Of potential concern is the replacement of seasonal wetlands with more permanent wetlands, which was apparent in nearly all of these studies, and appears to be a common outcome of wetland creation projects [130, 131]. The influence of biotic interactions on community structure tends to increase as water permanence increases [132–134]. Lengthening of wetland hydroperiod increases predation potential (e.g., through fish colonization), and in some cases promotes invasion by nonnative amphibians. For example, Fuller et al.  found that extended wetland hydroperiods due to the creation of side channels and mine tailing ponds along the Trinity River in California, USA, increased habitat suitability for the invasive bullfrog. Similarly, Maret et al.  concluded that replacement of seasonal marshes with permanent cattle tanks in Arizona negatively impacted endangered Sonoran tiger salamanders (Ambystoma tigrinum stebbinsi) by increasing invasions of fish and bullfrogs. Because hydroperiod dynamics exert such a strong influence on amphibian communities, we recommend that managers consider the surrounding wetland community when engaging in wetlands creation initiatives . Bedford  provided a conceptual background for this approach.
- D. B. Wake, “Declining amphibian populations,” Science, vol. 253, no. 5022, p. 860, 1991.
- J. H. K. Pechmann, D. E. Scott, R. D. Semlitsch, J. P. Caldwell, L. J. Vitt, and J. W. Gibbons, “Declining amphibian populations: the problem of separating human impacts from natural fluctuations,” Science, vol. 253, no. 5022, pp. 892–895, 1991.
- J. H. K. Pechmann and H. M. Wilbur, “Putting declining amphibian populations in perspective: natural fluctuations and human impacts,” Herpetologica, vol. 50, no. 1, pp. 65–84, 1994.
- R. A. Alford and S. J. Richards, “Global amphibian declines: a problem in applied ecology,” Annual Review of Ecology and Systematics, vol. 30, no. 1, pp. 133–165, 1999.
- T. J. C. Beebee and R. A. Griffiths, “The amphibian decline crisis: a watershed for conservation biology?” Biological Conservation, vol. 125, no. 3, pp. 271–285, 2005.
- S. N. Stuart, J. S. Chanson, N. A. Cox et al., “Status and trends of amphibian declines and extinctions worldwide,” Science, vol. 306, no. 5702, pp. 1783–1786, 2004.
- J. P. Collins and A. Storfer, “Global amphibian declines: sorting the hypotheses,” Diversity and Distributions, vol. 9, no. 2, pp. 89–98, 2003.
- R. M. Lehtinen, S. M. Galatowitsch, and J. R. Tester, “Consequences of habitat loss and fragmentation for wetland amphibian assemblages,” Wetlands, vol. 19, no. 1, pp. 1–12, 1999.
- J. P. Gibbs, “Wetland loss and biodiversity conservation,” Conservation Biology, vol. 14, no. 1, pp. 314–317, 2000.
- R. A. Alford, P. M. Dixon, and J. H. K. Pechmann, “Global amphibian population declines,” Nature, vol. 412, no. 6846, pp. 499–500, 2001.
- W. J. Mitsch and J. G. Gosselink, Wetlands, Wiley, New York, NY, USA, 4th edition, 2007.
- C. Scott Findlay and J. Houlahan, “Anthropogenic correlates of species richness in southeastern Ontario wetlands,” Conservation Biology, vol. 11, no. 4, pp. 1000–1009, 1997.
- R. D. Semlitsch, “Principles for management of aquatic-breeding amphibians,” Journal of Wildlife Management, vol. 64, no. 3, pp. 615–631, 2000.
- J. E. Houlahan and C. S. Findlay, “The effects of adjacent land use on wetland amphibian species richness and community composition,” Canadian Journal of Fisheries and Aquatic Sciences, vol. 60, no. 9, pp. 1078–1094, 2003.
- M. M. Brinson, “A hydrogeomorphic classification for wetlands,” Tech. Rep. WRP-DE-4, U.S. Army Corps of Engineers, Waterways Experiment Station, Vicksburg, Miss, USA, 1993.
- D. R. Smith, A. Ammann, C. Bartoldus, and M. M. Brinson, “An approach for assessing wetland functions using hydrogeomorphic classification, reference wetlands, and functional indices,” Tech. Rep. WRP-DE-9, U.S. Army Corps of Engineers, Waterways Experiment Station, Vicksburg, Miss, USA, 1995.
- C. V. Noble and L. Carpenter, “Hydrogeomorphic approach to assessing wetland functions: guidelines for developing regional guidebooks,” Tech. Rep. ERDC/EL TR-09-6, U.S. Army Corps of Engineers, Research and Development Center, Vicksburg, Miss, USA, 2009.
- R. F. Wilson and W. J. Mitsch, “Improving the success of wetland creation and restoration with know-how, time, and self-design,” Ecological Applications, vol. 6, no. 1, pp. 77–83, 1996.
- C. A. Cole, “HGM and wetland functional assessment: six degrees of separation from the data?” Ecological Indicators, vol. 6, no. 3, pp. 485–493, 2006.
- C. I. Kettlewell, V. Bouchard, D. Porej et al., “An assessment of wetland impacts and compensatory mitigation in the Cuyahoga river watershed, Ohio, USA,” Wetlands, vol. 28, no. 1, pp. 57–67, 2008.
- National Research Council, Compensating for Wetland Losses under the Clean Water Act, National Academy Press, Washington, DC, USA, 2001.
- M. M. Brinson and R. Rheinhardt, “The role of reference wetlands in functional assessment and mitigation,” Ecological Applications, vol. 6, no. 1, pp. 69–76, 1996.
- W. J. Mitsch and S. E. Jørgensen, Ecological Engineering and Ecosystem Restoration, Wiley, New York, NY, USA, 1st edition, 2004.
- S. E. G. Findlay, E. Kiviat, W. C. Nieder, and E. A. Blair, “Functional assessment of a reference wetland set as a tool for science, management and restoration,” Aquatic Sciences, vol. 64, no. 2, pp. 107–117, 2002.
- M. M. Brinson, “Classification of wetlands,” in Wetlands, B. A. LePage, Ed., pp. 95–113, Springer, Dordrecht, Netherlands, 2011.
- R. J. Stevenson and F. R. Hauer, “Integrating Hydrogeomorphic and Index of Biotic Integrity approaches for environmental assessment of wetlands,” Journal of the North American Benthological Society, vol. 21, no. 3, pp. 502–513, 2002.
- A. D. Jacobs, M. E. Kentula, and A. T. Herlihy, “Developing an index of wetland condition from ecological data: an example using HGM functional variables from the Nanticoke watershed, USA,” Ecological Indicators, vol. 10, no. 3, pp. 703–712, 2010.
- E. D. Stein, M. M. Brinson, M. C. Rains, W. Kleindl, and F. R. Hauer, “Wetland assessment alphabet soup: how to choose (or not choose) the right assessment method,” Wetland Science and Practice, vol. 26, no. 4, pp. 20–24, 2009.
- C. K. Balcombe, J. T. Anderson, R. H. Fortney, and W. S. Kordek, “Wildlife use of mitigation and reference wetlands in West Virginia,” Ecological Engineering, vol. 25, no. 1, pp. 85–99, 2005.
- D. Malakoff, “Restored wetlands flunk real-world test,” Science, vol. 280, no. 5362, pp. 371–372, 1998.
- J. B. Zedler and J. C. Callaway, “Tracking wetland restoration: do mitigation sites follow desired trajectories?” Restoration Ecology, vol. 7, no. 1, pp. 69–73, 1999.
- H. M. Wilbur, “Complex life cycles,” Annual Review of Ecology and Systematics, vol. 11, pp. 67–93, 1980.
- K. D. Wells, The Ecology & Behavior of Amphibians, University of Chicago Press, Chicago, Ill, USA, 2007.
- P. B. Pearman, “Effects of habitat size on tadpole populations,” Ecology, vol. 74, no. 7, pp. 1982–1991, 1993.
- K. J. Babbitt, “The relative importance of wetland size and hydroperiod for amphibians in southern New Hampshire, USA,” Wetlands Ecology and Management, vol. 13, no. 3, pp. 269–279, 2005.
- M. J. Baber and K. J. Babbitt, “The relative impacts of native and introduced predatory fish on a temporary wetland tadpole assemblage,” Oecologia, vol. 136, no. 2, pp. 289–295, 2003.
- C. A. Pearl, M. J. Adams, N. Leuthold, and R. B. Bury, “Amphibian occurrence and aquatic invaders in a changing landscape: implications for wetland mitigation in the Willamette Valley, Oregon, USA,” Wetlands, vol. 25, no. 1, pp. 76–88, 2005.
- R. Boyer and C. E. Grue, “The need for water quality criteria for frogs,” Environmental Health Perspectives, vol. 103, no. 4, pp. 352–357, 1995.
- K. Barrett, C. Guyer, and D. Watson, “Water from Urban streams slows growth and speeds metamorphosis in fowler's toad (Bufo fowleri) Larvae,” Journal of Herpetology, vol. 44, no. 2, pp. 297–300, 2010.
- K. J. Babbitt and G. W. Tanner, “Effects of cover and predator size on survival and development of Rana utricularia tadpoles,” Oecologia, vol. 114, no. 2, pp. 258–262, 1998.
- R. J. Monello and R. G. Wright, “Amphibian habitat preferences among artificial ponds in the Palouse region of northern Idaho,” Journal of Herpetology, vol. 33, no. 2, pp. 298–303, 1999.
- K. Kopp, M. Wachlevski, and P. C. Eterovick, “Environmental complexity reduces tadpole predation by water bugs,” Canadian Journal of Zoology, vol. 84, no. 1, pp. 136–140, 2006.
- T. Hartel, S. Nemes, D. Cogǎlniceanu et al., “The effect of fish and aquatic habitat complexity on amphibians,” Hydrobiologia, vol. 583, no. 1, pp. 173–182, 2007.
- C. R. Goldman, “Aquatic primary production,” Integrative and Comparative Biology, vol. 8, no. 1, pp. 31–42, 1968.
- D. B. Seale, “Influence of amphibian larvae on primary production, nutrient flux, and competition in a pond ecosystem,” Ecology, vol. 61, no. 6, pp. 1531–1550, 1980.
- J. D. Rouse, C. A. Bishop, and J. Struger, “Nitrogen pollution: an assessment of its threat to amphibian survival,” Environmental Health Perspectives, vol. 107, no. 10, pp. 799–803, 1999.
- G. S. Schuytema and A. V. Nebeker, “Effects of ammonium nitrate, sodium nitrate, and urea on red-legged frogs, pacific treefrogs, and African clawed frogs,” Bulletin of Environmental Contamination and Toxicology, vol. 63, no. 3, pp. 357–364, 1999.
- A. B. Sacerdote and R. B. King, “Dissolved oxygen requirements for hatching success of two ambystomatid salamanders in restored ephemeral ponds,” Wetlands, vol. 29, no. 4, pp. 1202–1213, 2009.
- P. T. J. Johnson, J. M. Chase, K. L. Dosch et al., “Aquatic eutrophication promotes pathogenic infection in amphibians,” Proceedings of the National Academy of Sciences of the United States of America, vol. 104, no. 40, pp. 15781–15786, 2007.
- D. Vasconcelos and A. J. K. Calhoun, “Monitoring created seasonal pools for functional success: a six-year case study of amphibian responses, Sears Island, Maine, USA,” Wetlands, vol. 26, no. 4, pp. 992–1003, 2006.
- B. Banks and T. J. C. Beebee, “Factors influencing breeding site choice by the pioneering amphibian Bufo calamita,” Holarctic Ecology, vol. 10, no. 1, pp. 14–21, 1987.
- Z. Wu, Y. Li, Y. Wang, and M. J. Adams, “Diet of introduced Bullfrogs (Rana catesbeiana): predation on and diet overlap with native frogs on Daishan Island, China,” Journal of Herpetology, vol. 39, no. 4, pp. 668–674, 2005.
- L. S. Vogel and J. H. K. Pechmann, “Response of fowler's toad (Anaxyrus fowleri) to competition and hydroperiod in the presence of the invasive coastal plain toad (Incilius nebulifer),” Journal of Herpetology, vol. 44, no. 3, pp. 382–389, 2010.
- J. W. Snodgrass, M. J. Komoroski, A. L. Bryan, and J. Burger, “Relationships among isolated wetland size, hydroperiod, and amphibian species richness: implications for wetland regulations,” Conservation Biology, vol. 14, no. 2, pp. 414–419, 2000.
- T. A. G. Rittenhouse and R. D. Semlitsch, “Distribution of amphibians in terrestrial habitat sorrounding wetlands,” Wetlands, vol. 27, no. 1, pp. 153–161, 2007.
- D. B. Means, “Vertebrate faunal diversity of longleaf pine ecosystems,” in The Longleaf Pine Ecosystem, S. Jose, E. J. Jokela, and D. L. Miller, Eds., pp. 157–213, Springer, New York, NY, USA, 2006.
- K. A. Berven, “Factors affecting population fluctuations in larval and adult stages of the wood frog (Rana sylvatica),” Ecology, vol. 71, no. 4, pp. 1599–1608, 1990.
- J. W. Petranka, C. K. Smith, and A. F. Scott, “Identifying the minimal demographic unit for monitoring pond-breeding amphibians,” Ecological Applications, vol. 14, no. 4, pp. 1065–1078, 2004.
- D. R. Church, “Role of current versus historical hydrology in amphibian species turnover within local pond communities,” Copeia, no. 1, pp. 115–125, 2008.
- J. W. Petranka and C. T. Holbrook, “Wetland restoration for amphibians: should local sites be designed to support metapopulations or patchy populations?” Restoration Ecology, vol. 14, no. 3, pp. 404–411, 2006.
- E. P. Ashley and J. T. Robinson, “Road mortality of amphibians, reptiles and other wildlife on the long point causeway, Lake Erie, Ontario,” Canadian Field-Naturalist, vol. 110, no. 3, pp. 403–412, 1996.
- M. J. Aresco, “Mitigation measures to reduce highway mortality of turtles and other herpetofauna at a north Florida lake,” Journal of Wildlife Management, vol. 69, no. 2, pp. 549–560, 2005.
- D. Florance, J. K. Webb, T. Dempster, M. R. Kearney, A. Worthing, and M. Letnic, “Excluding access to invasion hubs can contain the spread of an invasive vertebrate,” Proceedings of the Royal Society B, vol. 278, no. 1720, pp. 2900–2908, 2011.
- M. A. Gaston, A. Fuji, F. W. Weckerly, and M. R. J. Forstner, “Potential component allee effects and their impact on wetland management in the conservation of endangered anurans,” PLoS ONE, vol. 5, no. 4, Article ID e10102, 2010.
- M. J. Lacki, J. W. Hummer, and H. J. Webster, “Mine-drainage treatment wetland as habitat for herpetofaunal wildlife,” Environmental Management, vol. 16, no. 4, pp. 513–520, 1992.
- L. C. Batty, “The potential importance of mine sites for biodiversity,” Mine Water and the Environment, vol. 24, no. 2, pp. 101–103, 2005.
- R. H. Kadlec and S. Wallace, Treatment Wetlands, CRC Press, Boca Raton, Fla, USA, 2nd edition, 2009.
- J. M. R. Baker and T. R. Halliday, “Amphibian colonization of new ponds in an agricultural landscape,” Herpetological Journal, vol. 9, no. 2, pp. 55–63, 1999.
- K. J. Babbitt and G. W. Tanner, “Use of temporary wetlands by anurans in a hydrologically modified landscape,” Wetlands, vol. 20, no. 2, pp. 313–322, 2000.
- A. Chovanec, F. Schiemer, H. Waidbacher, and R. Spolwind, “Rehabilitation of a heavily modified river section of the Danube in Vienna (Austria): biological assessment of landscape linkages on different scales,” International Review of Hydrobiology, vol. 87, no. 2-3, pp. 183–195, 2002.
- D. Porej and T. E. Hetherington, “Designing wetlands for amphibians: the importance of predatory fish and shallow littoral zones in structuring of amphibian communities,” Wetlands Ecology and Management, vol. 13, no. 4, pp. 445–455, 2005.
- J. T. Julian, C. D. Snyder, and J. A. Young, “The use of artificial impoundments by two amphibian species in the Delaware Water Gap National Recreation Area,” Northeastern Naturalist, vol. 13, no. 4, pp. 459–468, 2006.
- J. W. Petranka, E. M. Harp, C. T. Holbrook, and J. A. Hamel, “Long-term persistence of amphibian populations in a restored wetland complex,” Biological Conservation, vol. 138, no. 3-4, pp. 371–380, 2007.
- C. D. Shulse, R. D. Semlitsch, K. M. Trauth, and A. D. Williams, “Influences of design and landscape placement parameters on amphibian abundance in constructed wetlands,” Wetlands, vol. 30, no. 5, pp. 915–928, 2010.
- P. Joly and O. Grolet, “Colonization dynamics of new ponds, and the age structure of colonizing alpine newts, Triturus alpestris,” Acta Oecologica, vol. 17, no. 6, pp. 599–608, 1996.
- T. J. C. Beebee, “Changes in dewpond numbers and amphibian diversity over 20 years on chalk downland in Sussex, England,” Biological Conservation, vol. 81, no. 3, pp. 215–219, 1997.
- R. Rannap, A. Lõhmus, and L. Briggs, “Restoring ponds for amphibians: a success story,” Hydrobiologia, vol. 634, no. 1, pp. 87–95, 2009.
- J. M. Cunningham, A. J. K. Calhoun, and W. E. Glanz, “Pond-breeding amphibian species richness and habitat selection in a beaver-modified landscape,” Journal of Wildlife Management, vol. 71, no. 8, pp. 2517–2526, 2007.
- T. E. Fuller, K. L. Pope, D. T. Ashton, and H. H. Welsh Jr., “Linking the distribution of an invasive amphibian (Rana catesbeiana) to habitat conditions in a managed river system in northern California,” Restoration Ecology, vol. 19, no. 201, pp. 204–213, 2011.
- J. S. Denton and T. J. C. Beebee, “Effects of predator interactions, prey palatability and habitat structure on survival of natterjack toad Bufo calamita larvae in replicated semi-natural ponds,” Ecography, vol. 20, no. 2, pp. 166–174, 1997.
- M. J. Adams, C. A. Pearl, and R. B. Bury, “Indirect facilitation of an anuran invasion by non-native fishes,” Ecology Letters, vol. 6, no. 4, pp. 343–351, 2003.
- E. E. Werner and M. A. McPeek, “Direct and indirect effects of predators on two anuran species along an environmental gradient,” Ecology, vol. 75, no. 5, pp. 1368–1382, 1994.
- L. M. Kurzava and P. J. Morin, “Tests of functional equivalence: complementary roles of salamanders and fish in community organization,” Ecology, vol. 79, no. 2, pp. 477–489, 1998.
- C. K. Adams, D. Saenz, and R. N. Conner, “Palatability of twelve species of anuran larvae in eastern Texas,” American Midland Naturalist, vol. 166, no. 1, pp. 211–223, 2011.
- R. D. Semlitsch and J. W. Gibbons, “Fish predation in size-structured populations of treefrog tadpoles,” Oecologia, vol. 75, no. 3, pp. 321–326, 1988.
- J. W. Petranka, C. A. Kennedy, and S. S. Murray, “Response of amphibians to restoration of a southern Appalachian wetland: a long-term analysis of community dynamics,” Wetlands, vol. 23, no. 4, pp. 1030–1042, 2003.
- J. A. Simon, J. W. Snodgrass, R. E. Casey, and D. W. Sparling, “Spatial correlates of amphibian use of constructed wetlands in an urban landscape,” Landscape Ecology, vol. 24, no. 3, pp. 361–373, 2009.
- A. B. Brand and J. W. Snodgrass, “Value of artificial habitats for amphibian reproduction in altered landscapes: contributed paper,” Conservation Biology, vol. 24, no. 1, pp. 295–301, 2010.
- D. Lesbarrères, M. S. Fowler, A. Pagano, and T. Lodé, “Recovery of anuran community diversity following habitat replacement,” Journal of Applied Ecology, vol. 47, no. 1, pp. 148–156, 2010.
- J. H. K. Pechmann, R. A. Estes, D. E. Scott, and J. W. Gibbons, “Amphibian colonization and use of ponds created for trial mitigation of wetland loss,” Wetlands, vol. 21, no. 1, pp. 93–111, 2001.
- J. A. Henning and G. Schirato, “Amphibian use of Chehalis River floodplain wetlands,” Northwestern Naturalis, vol. 87, no. 3, pp. 209–214, 2006.
- P. Galán, “Colonization of spoil benches of an opencast lignite mine in northwest Spain by amphibians and reptiles,” Biological Conservation, vol. 79, no. 2-3, pp. 187–195, 1997.
- C. E. Stevens, A. W. Diamond, and T. S. Gabor Shane, “Anuran call surveys on small wetlands in Prince Edward Island, Canada restored by dredging of sediments,” Wetlands, vol. 22, no. 1, pp. 90–99, 2002.
- R. M. Lehtinen and S. M. Galatowitsch, “Colonization of restored wetlands by amphibians in Minnesota,” American Midland Naturalist, vol. 145, no. 2, pp. 388–396, 2001.
- C. F. Bowers, H. G. Hanlin, D. C. Guynn, J. P. McLendon, and J. R. Davis, “Herpetofaunal and vegetational characterization of a thermally-impacted stream at the beginning of restoration,” Ecological Engineering, vol. 15, no. 1, pp. S101–S114, 2000.
- J. Colding, J. Lundberg, S. Lundberg, and E. Andersson, “Golf courses and wetland fauna,” Ecological Applications, vol. 19, no. 6, pp. 1481–1491, 2009.
- Y. F. Lee, Y. M. Kuo, Y. H. Lin, W. C. Chu, H. H. Wang, and S. H. Wu, “Composition, diversity, and spatial relationships of anurans following wetland restoration in a managed tropical forest,” Zoological Science, vol. 23, no. 10, pp. 883–891, 2006.
- J. W. Petranka, S. S. Murray, and C. A. Kennedy, “Responses of amphibians to restoration of a southern Appalachian wetland: perturbations confound post-restoration assessment,” Wetlands, vol. 23, no. 2, pp. 278–290, 2003.
- D. Vasconcelos and A. J. K. Calhoun, “Movement patterns of adult and juvenile Rana sylvatica (LeConte) and Ambystoma maculatum (Shaw) in three restored seasonal pools in maine,” Journal of Herpetology, vol. 38, no. 4, pp. 551–561, 2004.
- K. S. Mierzwa, “Wetland mitigation and amphibians: preliminary observations at a southwestern Illinois bottomland hardwood forest restoration site,” Journal of the Iowa Academy of Science, vol. 103, no. 3, pp. 191–194, 2000.
- M. Lannoo, Amphibian Declines: The Conservation Status of United States Species, University of California Press, Berkeley, Calif, USA, 2005.
- R. D. Bartlett and P. P. Bartlett, Guide and Reference to the Amphibians of Eastern and Central North America (North of Mexico), University Press of Florida, Tallahassee, Fla, USA, 2006.
- M. J. Mazerolle, M. Poulin, C. Lavoie, L. Rochefort, A. Desrochers, and B. Drolet, “Animal and vegetation patterns in natural and man-made bog pools: implications for restoration,” Freshwater Biology, vol. 51, no. 2, pp. 333–350, 2006.
- T. S. Nedland, A. Wolf, and T. Reed, “A reexamination of restored wetlands in Manitowoc County, Wisconsin,” Wetlands, vol. 27, no. 4, pp. 999–1015, 2007.
- J. G. Palis, “If you build it, they will come: herpetofaunal colonization of constructed wetlands and adjacent terrestrial habitat in the Cache River drainage of southern Illinois,” Transactions of the Illinois State Academy of Science, vol. 100, no. 2, pp. 177–189, 2007.
- E. D. Brodie Jr., D. R. Formanowicz Jr., and E. D. Brodie III, “The development of noxiousness of Bufo americanus tadpoles to aquatic insect predators,” Herpetologica, vol. 34, no. 3, pp. 302–306, 1978.
- E. D. Brodie Jr. and D. R. Formanowicz Jr., “Antipredator mechanisms of larval anurans: protection of palatable individuals,” Herpetologica, vol. 43, no. 3, pp. 369–373, 1987.
- M. S. Gunzburger and J. Travis, “Critical literature review of the evidence for unpalatability of amphibian eggs and larvae,” Journal of Herpetology, vol. 39, no. 4, pp. 547–571, 2005.
- J. W. Petranka, M. E. Hopey, B. T. Jennings, S. D. Baird, and S. J. Boone, “Breeding habitat segregation of wood frogs and American toads: the role of interspecific tadpole predation and adult choice,” Copeia, no. 3, pp. 691–697, 1994.
- J. P. Collins, “Intrapopulation variation in the body size at metamorphosis and timing of metamorphosis in the bullfrog, Rana catesbeiana,” Ecology, vol. 60, no. 4, pp. 738–749, 1979.
- A. H. Wright and A. A. Wright, Handbook of Frogs and Toads of the United States and Canada, Comstock Publishing Company, Ithaca, NY, USA, 1949.
- R. N. Homan, B. S. Windmiller, and J. M. Reed, “Critical thresholds associated with habitat loss for two vernal pool-breeding amphibians,” Ecological Applications, vol. 14, no. 5, pp. 1547–1553, 2004.
- S. M. Blomquist and M. L. Hunter Jr., “A multi-scale assessment of amphibian habitat selection: wood frog response to timber harvesting,” Ecoscience, vol. 17, no. 3, pp. 251–264, 2010.
- T. A. Touré and G. A. Middendorf, “Colonization of herpetofauna to a created wetland,” Bulletin of the Maryland Herpetological Society, vol. 38, no. 4, pp. 99–117, 2002.
- L. P. Shoo, D. H. Olson, S. K. McMenamin, et al., “Engineering a future for amphibians under climate change,” Journal of Applied Ecology, vol. 48, no. 2, pp. 487–492, 2011.
- T. J. Maret, J. D. Snyder, and J. P. Collins, “Altered drying regime controls distribution of endangered salamanders and introduced predators,” Biological Conservation, vol. 127, no. 2, pp. 129–138, 2006.
- W. S. Hou, Y. H. Chang, H. W. Wang, and Y. C. Tan, “Using the behavior of seven amphibian species for the design of banks of irrigation and drainage systems in Taiwan,” Irrigation and Drainage, vol. 59, no. 5, pp. 493–505, 2010.
- L. R. Gamble, K. McGarigal, C. L. Jenkins, and B. C. Timm, “Limitations of regulated “buffer zones” for the conservation of marbled salamanders,” Wetlands, vol. 26, no. 2, pp. 298–306, 2006.
- R. D. Semlitsch, “Biological delineation of terrestrial buffer zones for pond-breeding salamanders,” Conservation Biology, vol. 12, no. 5, pp. 1113–1119, 1998.
- R. D. Semlitsch and J. R. Bodie, “Biological criteria for buffer zones around wetlands and riparian habitats for amphibians and reptiles,” Conservation Biology, vol. 17, no. 5, pp. 1219–1228, 2003.
- D. Vasconcelos and A. J. K. Calhoun, “Movement patterns of adult and juvenile Rana sylvatica (LeConte) and Ambystoma maculatum (Shaw) in three restored seasonal pools in Maine,” Journal of Herpetology, vol. 38, no. 4, pp. 551–561, 2004.
- C. D. Shulse, R. D. Semlitsch, and K. M. Trauth, “Development of amphibian biotic index to evaluate wetland health in northern Missouri,” in Proceedings of the World Environmental and Water Resources Congress, S. Starrett, Ed., pp. 2657–2667, American Society of Civil Engineers, Reston, Va, USA, 2009.
- C. G. Becker, C. R. Fonseca, C. F. B. Haddad, R. F. Batista, and P. I. Prado, “Habitat split and the global decline of amphibians,” Science, vol. 318, no. 5857, pp. 1775–1777, 2007.
- C. J. Briggs, V. T. Vredenburg, R. A. Knapp, and L. J. Rachowicz, “Investigating the population-level effects of chytridiomycosis: an emerging infectious disease of amphibians,” Ecology, vol. 86, no. 12, pp. 3149–3159, 2005.
- K. G. Smith, K. R. Lips, and J. M. Chase, “Selecting for extinction: nonrandom disease-associated extinction homogenizes amphibian biotas,” Ecology Letters, vol. 12, no. 10, pp. 1069–1078, 2009.
- D. B. Means and P. E. Moler, “The pine barrens treefrog: fire, seepage bogs, and management implications,” in Technical Bulletin WL4, R. R. Odum and L. Landers, Eds., pp. 77–83, Georgia Game and Fish Division, Atlanta, Ga, USA, 1979.
- H. H. Welsh Jr., “Relictual amphibians and old-growth forests,” Conservation Biology, vol. 4, no. 3, pp. 309–319, 1990.
- A. L. Gallant, R. W. Klaver, G. S. Casper, and M. J. Lannoo, “Global rates of habitat loss and implications for amphibian conservation,” Copeia, no. 4, pp. 967–979, 2007.
- D. M. Marsh and P. C. Trenham, “Metapopulation dynamics and amphibian conservation,” Conservation Biology, vol. 15, no. 1, pp. 40–49, 2001.
- C. A. Cole and R. P. Brooks, “A comparison of the hydrologic characteristics of natural and created mainstem floodplain wetlands in Pennsylvania,” Ecological Engineering, vol. 14, no. 3, pp. 221–231, 2000.
- D. L. Gamble and W. J. Mitsch, “Hydroperiods of created and natural vernal pools in central Ohio: a comparison of depth and duration of inundation,” Wetlands Ecology and Management, vol. 17, no. 4, pp. 385–395, 2009.
- G. A. Wellborn, D. K. Skelly, and E. E. Werner, “Mechanisms creating community structure across a freshwater habitat gradient,” Annual Review of Ecology and Systematics, vol. 27, pp. 337–363, 1996.
- L. De Meester, S. Declerck, R. Stoks et al., “Ponds and pools as model systems in conservation biology, ecology and evolutionary biology,” Aquatic Conservation: Marine and Freshwater Ecosystems, vol. 15, no. 6, pp. 715–725, 2005.
- T. L. Tarr, M. J. Baber, and K. J. Babbitt, “Macroinvertebrate community structure across a wetland hydroperiod gradient in southern New Hampshire, USA,” Wetlands Ecology and Management, vol. 13, no. 3, pp. 321–334, 2005.
- T. J. Maret, J. D. Snyder, and J. P. Collins, “Altered drying regime controls distribution of endangered salamanders and introduced predators,” Biological Conservation, vol. 127, no. 2, pp. 129–138, 2006.
- A. J. K. Calhoun, N. A. Miller, and M. W. Klemens, “Conserving pool-breeding amphibians in human-dominated landscapes through local implementation of Best Development Practices,” Wetlands Ecology and Management, vol. 13, no. 3, pp. 291–304, 2005.
- B. L. Bedford, “The need to define hydrologic equivalence at the landscape scale for freshwater wetland mitigation,” Ecological Applications, vol. 6, no. 1, pp. 57–68, 1996.
- M. B. Dietsch, H. G. Hanlin, E. D. Jones, and L. D. Wike, “Dynamics of a herpetofaunal community in a restored freshwater wetland,” Bulletin of the Ecological Society of America, vol. 77, p. 113, 1996.
- S. Juni and C. R. Berry, “A biodiversity assessment of compensatory mitigation wetlands in eastern South Dakota,” Proceedings of the South Dakota Academy of Science, vol. 80, pp. 185–200, 2001.
- C. A. Pearl and J. Bowerman, “Observations of rapid colonization of constructed ponds by western toads (Bufo boreas) in Oregon, USA,” Western North American Naturalist, vol. 66, no. 3, pp. 397–401, 2006.