Habitats of the British amphibians (4): Agricultural lowlands and a general discussion of requirements

Ponds on about 80 km² of farmland in Sussex were investigated with regard to breeding amphibians. The survey covered five different geological strata and included 60 ponds (out of a total of more than 200) marked on 1:25000 ordnance maps. Eighteen of these ponds were found on inspection not to exist; the remainder were characterised with respect to surface area, depth, pH, ionic score, aquatic vegetation, fish fauna and surrounding terrestrial habitats as well as for amphibian species. 14 pools contained amphibians; Triturus vulgaris was the most widespread (9 ponds) closely followed by T. helveticus (8 ponds). T. cristatus occurred at 5 sites, Rana temporaria at 2 and Bufo bufo not at all. The 2 frog sites were very shallow pools on pasture/woodland interfaces; the ideal newt pond had no fish, was 0·5–1·0 m deep and < 200 m² in surface area with 5–50% aquatic vegetation cover and a low ionic score (< 400 parts/10⁶). Callitriche-containing ponds were particularly favoured, and with terrestrial habitat there was a requirement for a scrub or woodland component.     

Terrestrial activity and conservation of adult California red-legged frogs Rana aurora draytonii in coastal forests and grasslands

The federally threatened California red-legged frog Rana aurora draytonii occupies both aquatic and terrestrial habitats in its adult life stage. The terrestrial activities of this species are not well known and require documentation to assist in the development of appropriate levels of protection under the US Endangered Species Act. We studied the terrestrial activities of radio-tagged red-legged frogs (n=8-26) inhabiting a coastal watershed in Santa Cruz County, California, during 1997-1998. In particular, we investigated (1) the use of terrestrial habitats by non-migrating adults in relation to season, breeding chronology, and precipitation, and (2) adult migration behavior, including seasonal timing, duration, distances traveled, and the use of corridors. Non-migrating red-legged frogs occupied terrestrial habitats briefly (median=4-6 days) following infrequent summer rains, but resided nearly continuously on land (median=20-30 days) from the onset of the winter wet-season until breeding activities commenced 1-2 months later. All of the non-migrating frogs remained within 130 m of their aquatic site of residence (median <25 m). Intervals spent on land were again brief during mid/late winter (median=1-4 days), despite frequent and copious rainfall. Adult migration to and from breeding sites occurred from late October through mid-May (wet season). We monitored 25 migration events between aquatic sites that were 200-2800 m apart. Short distance movements (<300 m) were completed in 1-3 days, longer movements required up to 2 months. Most migrating frogs moved overland in approximately straight lines to target sites without apparent regard to vegetation type or topography. Riparian corridors were neither essential nor preferred as migration routes. Frogs traveling overland occurred in upland habitats as far as 500 m from water. Approximately 11-22% of the adult population was estimated to migrate to and from breeding sites annually, whereas the bulk of the adult population was resident at these sites. Adequate protection of red-legged frog populations inhabiting relatively undeveloped landscapes is liable to be achieved by retaining an array of suitable habitat elements within at least 100 m of occupied aquatic sites, and by imposing seasonal limitations on detrimental human activities occurring within this zone. Specific protections for migrating frogs are probably unwarranted in forest and rangeland environments because dispersal habitat is ubiquitous and migrating frogs are widely distributed across the landscape in space and time.     

Using habitat models to determine conservation priorities for pond-breeding amphibians in a privately-owned landscape of northern Idaho, USA

As rural landscapes experience increasing levels of residential development, the persistence of species that rely on them will depend on informed planning and management decisions. An understanding of habitat requirements is essential for setting priorities and developing landscape-level plans for the survival of these species. In many temperate rural landscapes, artificially created ponds may be the only wetlands available for aquatic reproduction by amphibians. The introduction of non-native fish into these ponds reduces survival and prohibits successful reproduction of many native pond-breeding amphibians. We surveyed 105 randomly selected wetlands in a primarily privately-owned, rural landscape in north Idaho, USA, for pond-breeding amphibian larvae in 2004 and 2005. We used an information theoretic multimodel inference and an algorithmic (random forests) approach to model habitat for each species based on local and landscape characteristics. We also used a mail survey to quantify how landowners value fish in their wetlands and their plans for future wetland development and fish stocking. Sixty-seven percent of pond owners reported that fishing in their pond was at least slightly important to them and 36% of owners indicated that they were at least 50% likely to add fish to their ponds in the next 5-10 years. Landscape change predictions for this area indicate that forests will become more open due to thinning; habitat models indicated that this is likely to be detrimental to long-toed salamanders and beneficial to Pacific treefrogs. Habitat models also indicate that Columbia spotted frog breeding sites consist of wetlands on flat ground with high solar insolation and that this species is sensitive to nearby development, indicating that as this landscape becomes further developed, this species may require habitat protection for persistence.     

The influence of acidic pond water on amphibians: A review

Acidic pond water may influence the reproduction of amphibians by causing direct mortality of embryos and larvae, and/or by disrupting trophic relationships between amphibians and other aquatic organisms. The embryo is the most sensitive stage of development and may abort soon after exposure to very low pH or eventually fail to hatch at a more moderate pH. Larvae are killed by disruption of Na and Cl balance. The toxicity of pond water is governed by complex interactions of pH, temperature, and the concentrations of Al, Ca, and organic acids. The reduction in recruitment associated with acidification of ponds affects both the local distribution and abundance of amphibians. The relative importance of different mechanisms of acidification of amphibian breeding sites is unknown. Amphibian breeding ponds are generally small, low in buffering capacity, and darkly stained by humic substances, making it difficult to discriminate between the contribution of acid from atmospheric input and natural sources (such as sphagnum and organic acids). It is however quite clear that temporary ponds are extremely vulnerable to acidification by rain and that they require continued monitoring in order to detect future changes in biological and/or chemical characteristics.     

Amphibian ecology and conservation in the urbanising world: A review

Urbanisation currently threatens over one-third of the world’s known amphibian species. The main threats of urbanisation to amphibian populations are habitat loss, habitat fragmentation and isolation, and degradation of habitat quality. A complex array of interacting biotic and abiotic factors impact amphibians in urban and urbanising landscapes. These can lead to a decrease in species richness and the abundance of individual species towards the centre of cities and towns. The ability of amphibians to disperse can be significantly reduced in urban and suburban landscapes. However, different species exhibit markedly different responses to urbanisation. Amphibian species that are habitat generalists or have relatively low dispersal requirements appear to be better able to survive in urban and suburban landscapes. There is insufficient information on the ecology of amphibians in urban and suburban areas, particularly in the tropics and sub-tropics, despite worldwide declines reported over past decades. Future research of amphibians in urban and suburban landscapes would greatly benefit by using long-term studies at sites along urban-rural gradients, conducted at both local and landscape scales. Research needs to be directed to the developing world in the tropics and sub-tropics, which has the highest rates of urbanisation. Research into amphibian ecology and conservation in the urbanising world would be improved through experimental approaches to determine the proximate causes of species’ responses to human modification of the landscape. Maintaining viable populations of amphibians in urban and suburban landscapes will require conservation strategies that consider key urbanisation processes (i.e. habitat availability and habitat quality) and the key responses and adaptations to urbanisation (i.e. species availability and species response). Conservation strategies for amphibians in urban and suburban landscapes need to include actions to prevent further loss and degradation of both terrestrial and aquatic habitat, and to reconnect the landscape to facilitate dispersal and long-term regional persistence of amphibian populations and communities.     

Influence of prey variables, food supply, and river restoration on the foraging activity of Daubenton’s bat (Myotis daubentonii) in the Shibetsu River, a large lowland river in Japan

To conserve the foraging habitat of Daubenton’s bat (Myotis daubentonii) in a large lowland river, we investigated the influence on this bat of prey variables (number or biomass) and insect origin (aquatic or terrestrial). We tested the hypothesis that river restoration (re-meandering) conducted in the Shibetsu River, northern Japan, enhances foraging habitat quality by increasing the abundance of aquatic insects. From June to September 2004, flying insects were collected using Malaise traps in restored and channelised reaches in the Shibetsu River. Bat activity was recorded by bat detectors placed near the Malaise traps in each of the two reaches. Foraging activity of Daubenton’s bat was more strongly related to the number of insects than to biomass, and to adult aquatic insects than to terrestrial insects. The high dependence of Daubenton’s bat on aquatic prey was attributed to the fact that aquatic insect numbers were always higher than those of terrestrial insects. Contrary to the hypothesis, Daubenton’s bat was more active in the channelised reach than the restored reach in all months except June, and it synchronized its foraging activity with the seasonal distribution of adult aquatic insects. However, the study was undertaken just two years after restoration and the riparian vegetation had not yet established itself. Our results demonstrate the importance of aquatic insect abundance for Daubenton’s bat throughout the seasons in large lowland rivers. A further decrease in aquatic insects, associated with progressive anthropogenic alteration of river environments, undoubtedly exerts a harmful influence on the conservation of this species.     

Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

Conservation planning for connectivity across marine, freshwater, and terrestrial realms

Conservation plans are usually developed for regions that encompass only one environmental realm (terrestrial, freshwater or marine) because of logistical, institutional and political constraints. This is inadequate because these realms often interact through processes that form, utilize and maintain interfaces or connections, which are essential for the persistence of some species and ecosystem functions. We present a conceptual framework for systematic conservation prioritization that explicitly accounts for the connectivity between the terrestrial, marine, and freshwater realms. We propose a classification of this connectivity that encompasses: (1) narrow interfaces, such as riparian strips; (2) broad interfaces, such as estuaries; (3) constrained connections, such as corridors of native vegetation used by amphibians to move between natal ponds and adult habitat; and (4) diffuse connections, such as the movements of animals between breeding and feeding habitats. We use this taxonomy of inter-realm connectivity to describe existing and new spatial conservation prioritization techniques that aim to promote the persistence of processes that operate between realms.     

Beaver (Castor canadensis) as a surrogate species for conserving anuran amphibians on boreal streams in Alberta, Canada

We explored the use of beaver (Castor canadensis) as a surrogate species for amphibian conservation on small (1st-4th-order) streams in the Boreal Foothills of west-central Alberta. Anuran call surveys indicated that beaver create breeding habitat for the boreal chorus frog (Pseudacris maculata), wood frog (Rana sylvatica) and western toad (Bufo boreas). No calling males of any species were recorded on unobstructed streams. Wood frog, the most abundant species, exhibited high rates of juvenile recruitment on beaver ponds. Pitfall traps captured more wood frogs on beaver ponds versus unobstructed streams, and most individuals (84%) were young-of-year. Abundance of young-of-year was strongly correlated with percent landscape occupied by beaver ponds indicating that anurans captured along streams originated in beaver ponds. Based on a novel combination of a digital elevation model and aerial photographs examined with GIS, statistical models showed that the probability of beaver pond occurrence on streams was positively associated with stream order and dependent on the interacting effects of distance to nearest forestry cutblock and availability of beaver foods (Populus spp.). We propose that the distribution and abundance of beaver ponds could be determined over large areas quickly and inexpensively by remote sensing and used to identify and monitor amphibian habitat, and possibly, populations. This work establishes the pre-eminence of beaver-created wetlands as amphibian habitat in the Boreal Foothills and that the incorporation of dam-building patterns into forest management strategies could aid amphibian conservation.     

The ecology of extinction: population fluctuation and decline in amphibians

Even among widespread species with high reproductive potentials and significant dispersal abilities, the probability of extinctions should be correlated both with population size variance and with the extent of population isolation. To address how variation in demographic characteristics and habitat requirements may reflect on the comparative risk of species decline, I examined 617 time series of population census data derived from 89 amphibian species using the normalized estimate of the realized rate of increase, ΔN, and its variance. Amphibians are demonstrably in general decline and exhibit a great range of dispersal abilities, demographic characteristics, and population sizes. I compared species according to life-history characteristics and habitat use. Among the populations examined, census declines outnumbered increases yet the average magnitudes for both declines and increases were not demonstrably different, substantiating findings of amphibian decline. This gives no support for the idea that amphibian population sizes are dictated by regimes featuring relatively rare years of high recruitment offset by intervening years of gradual decline such that declines may outnumber increases without negative effect. For any given population size, those populations living in large streams or in ponds had significantly higher variance than did populations of completely terrestrial or other stream-dwelling amphibians. This could not be related to life-history complexity as all the stream-breeding species examined have larvae and all of the wholly terrestrial species have direct development without a larval stage. Variance in ΔN was highest amongst the smallest populations in each comparison group. Estimated local extinction rates averaged 3.1% among pond-breeding frogs, 2.2% for pond-breeding salamanders, and negligible for both stream-breeding and terrestrial direct-developing species. Recoveries slightly exceeded extinctions among European pond-breeding frogs but not among North American pond-breeding frogs. Less common species had greater negative disparities between extinctions and recoveries. Species with highly fluctuating populations and high frequencies of local extinctions living in changeable environments, such pond- and torrent-breeding amphibians, may be especially susceptible to curtailment of dispersal and restriction of habitat.     

Long term monitoring of amphibian populations with respect to the effects of acidic deposition

Amphibians breed in a variety of aquatic habitats in the United States. While the tolerance of low pH has been examined for many species of amphibians, those that breed in temporary ponds have been the most studied in terms of acidic precipitation. Some of the latter have been shown to have lethal limits for low pH that are close to or above the pH's found in many breeding ponds. Temporary ponds occur in large numbers in the areas of the country most affected by acidic precipitation. Many of these ponds are susceptible to acidification and reduced reproductive success of sensitive species of salamanders has been observed in acidic ponds. Additional practical and logistical considerations clearly make temporary ponds, and certain salamanders that breed in them, the best candidates for monitoring the long term effects of acidic precipitation on amphibian populations. We recommend a cost efficient monitoring scheme across three categories of acidic temporary ponds (low, medium, and high) that incorporates (a) chemical analyses of pond samples, (b) a census of the annual deposition of egg masses, as well as assessment of embryonic mortality in enclosed and unenclosed egg masses in the ponds, and (c) sampling of larval success. This plan offers the best opportunity for detecting changes in amphibian populations that are correlated with the acidity of precipitation, and would provide desperately needed long term data on the abundance of these amphibians.     

Habitat selection by amphibians across an agricultural land-heathland transect in Britain

Potential and actual breeding sites for amphibians have been investigated in an area encompassing agricultural land and uncultivated heathland. pH and conductivity measurements have been made together with species records and some other aspects of habitat structure noted. Ponds around the farmland-heathland interface were especially well used by amphibians; there had been major losses of ponds in the agricultural zone, while those further out on the heathland were usually highly oligotrophic and capable of supporting only Triturus helveticus at most. Populations of the newts T. helveticus and T. vulgaris were composed of significantly smaller individuals in heathland ponds than in other sites. Implications of this study for conservation are discussed.     

Assessing threats to pool-breeding amphibian habitat in an urbanizing landscape

Geographically-based threat assessments are important for identifying natural resources at risk, yet have rarely been applied to identify habitat conservation priorities for imperiled organisms at a local scale. Pool-breeding amphibians have complex life cycles that place them at risk from habitat loss and fragmentation both in wetlands and in adjacent uplands. Because the most rapidly growing cause of habitat degradation in North America has been urbanization, a threat analysis of pool-breeding amphibian habitat should both be dynamic, i.e., sensitive to land-use change, and comprehensive, recognizing traditional protected area networks as well as less formal conservation assets (e.g., land-use regulations). To assess threats to wood frogs (Rana sylvatica) and spotted salamanders (Ambystoma maculatum) in a rapidly urbanizing, forested region of New England (USA) we examined gaps in the current protection network, as well as changing human settlement patterns. We found that greater than 50% of 542 potential breeding pools delineated using low-level infra-red aerial photography (median area 379.5 m²) were not represented on National Wetland Inventory (U.S.FWS) maps, and thus de facto at risk. Most importantly, conservation lands and regulatory protections failed to protect 46% of potential breeding pools and 80% of adjacent non-breeding habitat. While an assessment of human settlement patterns projected that only 5% of the region contained high quality amphibian habitat under acute development pressure, nearly half of the region (44.7%) had attained a moderate threat level, highlighting the importance of conservation planning during early stages of urbanization. We conclude by illustrating the role for multiple conservation strategies when protecting functional landscapes for pool-breeding amphibians.     

Core terrestrial habitat for conservation of local populations of salamanders and wood frogs in agricultural landscapes

Pond-breeding amphibians require aquatic and terrestrial habitats to complete their lifecycles, and preservation of both habitats is necessary for maintaining local populations. Current wetland regulations focus primarily on aquatic habitats, and criteria to define critical upland habitats and regulations to protect them are often ambiguous or lacking. We examined the association between the presence of seven pond-breeding amphibian species and the landscape composition surrounding 54 wetlands located within the Till Plains and the Glaciated Plateau ecoregions of Ohio, USA. We quantified landscape composition within 200 m of the wetland (“core terrestrial zone”) and the area extending from 200 m to 1 km from the wetland (“broader landscape context zone”). We constructed binary logistic regression models for each species, and evaluated them using Akaike Information Criterion. Presence of spotted salamanders (Ambystoma maculatum), Jefferson's salamander complex (A. jeffersonianum) and smallmouth salamanders (A. texanum) was positively associated with the amount of forest within the core zone. Presence of wood frogs (Rana sylvatica) was positively associated with the amount of forest within the core zone and the amount of forest within the broader landscape context zone. Presence of tiger salamanders (A. tigrinum tigrinum) was negatively associated with the cumulative length of paved roads within 1 km of the site, and presence of red-spotted newts (Notophthalmus v. viridescens) was negatively associated with the average linear distance to the five nearest wetlands. Overall salamander diversity was positively associated with the amount of forest within the core zone, and negatively associated with the presence of predatory fish and cumulative length of paved roads within 1 km of the site. Our results confirm the strong association between the structure of surrounding upland areas and amphibian diversity at breeding ponds, and stress the importance of preserving core terrestrial habitat around wetlands for maintaining amphibian diversity.     

Multi-scale effect of landscape processes and habitat quality on newt abundance: Implications for conservation

Recent studies in population dynamics suggest that landscape processes and habitat quality act at different scales on population abundances, but very few have modelled their simultaneous effects. However, at a time of large declines in natural populations, it is essential to understand such multivariate components. We tested the hypothesis that natural populations of palmate newts (Triturus helveticus) are affected on three scales: breeding patch (pond), habitat complementation (terrestrial cover), and metapopulation structure (density of ponds, surrounding populations). We conducted our survey in 130 ponds from southern France (Larzac) and analysed data with generalized additive models (GAM). Two main novel results emerge from these models: (1) the three landscape scales have significant effects on newt abundance, with more newts in deep, vegetated ponds, devoid of fish and surrounded by wooded areas and inhabited ponds; (2) the quality of the surrounding breeding patches is of primary importance in determining the abundance at core sites in a complex way: high abundances are associated positively with high densities of inhabited ponds, but negatively with the number of surrounding ponds. Deforestation, invasive species and abandonment of ponds all have negative impacts on the persistence of palmate newt populations. Future studies should encompass landscapes at different scales and incorporate the habitat quality in surrounding sites to better understand population dynamics and provide adequate conservation measures.     

Evaluation of translocation as a tool for mitigating development threats to great crested newts (Triturus cristatus) in England, 1990-2001

Great crested newts (Triturus cristatus) are protected under European and UK legislation, but are frequently the subject of conflict between development and conservation in England. When this occurs, the developer is legally obliged to instigate a mitigation plan to reduce the impacts on the newts. This usually involves the translocation of newts coupled with habitat enhancement and creation. We reviewed mitigation projects carried out in England between 1990 and 2001 by (1) analysing licensing information collected by the governmental licensing authorities; and (2) a questionnaire survey of a sample of mitigation projects. Over half of the licensed projects on file contained no report of the work undertaken. There was an increase in the number of new translocation projects from less than 10 a year in the early 1990s to over 80 a year by 2000. This translates into about £1.5 million per year currently being spent on great crested newt mitigation projects. Most of these projects involved in situ translocations of newts to areas within or adjacent to the development site. The number of newts translocated per project declined over the same period, and was related to the total area of habitat destroyed and work effort. About 27% of great crested newt terrestrial habitat was destroyed during the developments along with about half of all ponds. Although the number of new ponds created compensated for the number of known great crested newt ponds lost, there was a net loss in terms of overall area of aquatic habitat. Where follow-up monitoring of translocations was conducted, there was evidence of breeding at most sites one-year post-development, but it is unclear whether these populations were sustainable in the long-term.     

Ranging behaviour of greater flamingos during the breeding and post-breeding periods: Linking connectivity to biological processes

Connectivity among habitat patches has mainly been studied in relation to species-specific behaviours. However, the movements of animals among patches may have different functions, and tolerance to gaps between habitat patches may vary between these functions. We tracked the movements of greater flamingos during the breeding and post-breeding periods with the aim of illustrating how the degree of connectivity may vary depending on the biological processes underlying the movements between wetlands. Most foraging sites used by breeding adults in a colony in southern Spain were within 200 km of the colony site, although some birds eventually moved 400 km. After the breeding season, the adults remained for several weeks in specific wetlands, moving to other sites located 280-2100 km away to overwinter. During these movements the birds may use stopover sites, the conservation of which may be critical to facilitate long-range movements. Our results suggest that wetland connectivity during chick rearing does not seem to be determined by whether or not central-place foraging flamingos are able to reach wetlands located at the longest distance that they are able to fly during non-stop flights (>1000 km), but by whether they are able to sustain the energetic costs derived from frequent commuting. In contrast, long-distance movements were occasionally undertaken during the post-breeding period. The energetic costs of such flights could be paid by flamingos because these movements were infrequent. Thus, in the case of flamingos, connectedness thresholds between wetlands vary depending on the biological process involved (chick rearing or post-breeding movements). This emphasizes the need to consider different types of connectivity in conservation planning.     

Grasslands as movement barriers for a forest-associated salamander: Migration behavior of adult and juvenile salamanders at a distinct habitat edge

The persistence of amphibian populations in fragmented landscapes requires dispersal and recolonization of habitat patches after local extinction. These processes entail individuals crossing habitat edges. Edge permeability integrates the behavior of individuals with the vegetative structure of the habitat edge and may influence the dispersal rates between habitat patches. We used drift fences, radio telemetry, and an experimental displacement to examine the movement behavior of juvenile and adult spotted salamanders (Ambystoma maculatum) at a pond located in continuous forest and a pond located on a distinct forest-grassland edge. At the pond on the habitat edge, adult salamanders migrated to and from the forested side of the pond. Resident adults with transmitters migrated to forested habitat without approaching the habitat edge. Displaced adults with transmitters halted emigration movements when they approached the habitat edge. None of the radio-tagged adults were observed more than a few meters into the grassland. Recently metamorphosed juveniles exiting the grassland side of the pond changed their direction of movement and migrated into the forested habitat. We conclude that salamanders oriented movements with respect to features in the terrestrial habitat, detected the habitat edge, and behaviorally avoided the grassland. Exploring the permeability of habitat edges will improve our understanding of dispersal within fragmented landscapes and enhance efforts to conserve regional populations of amphibians.     

Amphibian production in forested landscapes in relation to wetland hydroperiod: A case study of vernal pools and beaver ponds

Many amphibian species are plastic in their selection of breeding sites and use both short- and longer-hydroperiod wetlands. Understanding which wetland types are most important to amphibian production is critical for focusing conservation efforts. We compared reproduction of wood frogs (Rana sylvatica) and spotted salamanders (Ambystoma maculatum) and production of wood frog juveniles between seasonal (vernal pools) and semi-permanent (beaver-created ponds) wetlands in the central Adirondack region of the northeastern United States. In 65 seasonal wetlands and 37 semi-permanent wetlands over 3 years, densities of wood frog egg masses were similar, but seasonal wetlands contained 4× as many spotted salamander egg masses as did semi-permanent wetlands. For wood frogs, survival to metamorphosis and juvenile production were an order of magnitude higher in semi-permanent wetlands. Models of wood frog production linked to the climatic record predicted that, depending on annual variation in rainfall and survival to metamorphosis, semi-permanent wetlands may produce annually 1.2–23× the number of wood frogs that seasonal wetlands produce within the study area. Models incorporating variation in demographic traits throughout the range of wood frogs suggest that higher survival to metamorphosis in beaver ponds and higher clutch densities in vernal pools strongly influence metamorph production on a regional basis. Semi-permanent wetlands make major contributions to amphibian production in our region and may also be an important source of colonists to seasonal wetlands following periods of drought. Protection of wetlands along the landscape-level continuum of hydroperiod therefore may be critical for maintaining and restoring amphibian populations.