Carbon dioxide efflux and pCO2 in soils of threeQuercus ilex montane forests

Soil CO₂ efflux and pCO₂ in the soil atmosphere were measured during one year at three montane sites of Mediterranean sclerophyllous forests in NE Spain. Two sites were located in the upper and lower slopes of a small catchment in the Prades mountains (mean precipitation 550 mm year^–1), and a third site was located on a lower slope in the Montseny mountains (mean precipitation 900 mm year^–1). The three sites were similar in bedrock and vegetation, but differed in soil characteristics and water availability. Seasonal variation of CO₂ efflux and soil pCO₂ were affected by soil temperature and, to a lesser extent, by soil moisture. Annual mean soil CO₂ efflux (considered as soil respiration) was similar at Montseny and at the comparably located site at Prades (83 ± 18 S.E. vs. 75 ± 9 mg CO₂ m^–2 hour^–1 , respectively), and was highest at the Prades upper slope site (122 ± 22 mg C0₂ m^–2 hour^–1 ). Despite those relatively similar CO₂ effluxes, mean soil pCO₂ was much higher at both Prades sites than at Montseny. Soil pCO₂ always increased with depth at Prades while maxima pCO₂ at Montseny were often at 20–30 cm depth. A model based on gas diffusion theory was able to explain why soil pCO₂ was much higher at Prades than at Montseny, and to reproduce the shape of the vertical profile of pCO₂ at the Prades soils. Nevertheless, the model failed to simulate the soil pCO₂ maximum found at 20–30 cm depth at the Montseny site. Model simulations using a time-variable CO₂ production rate suggested that pCO₂ maxima at intermediate depth could be the result of a transient situation instead of an equilibrium one.     

Effects of an introduced crayfish, Procambarus clarkii, on the distribution of south-western Iberian amphibians in their breeding habitats

The introduction of exotic species in aquatic habitats is one of the causes for the amphibian declines observed worldwide. In the 1970s, the red swamp crayfish Procambarus clarkii was introduced in the southwest Iberian Peninsula, where no native crayfish occur. In this study we assess the effect of P. clarkii presence in the breeding site distribution of each of the 13 southwest Iberian amphibians, while simultaneously accounting for the effects of potentially confounding habitat variables, as well as for the effects of the other large aquatic predators in the study area – predatory fish. Amphibian species richness was lower in places where P. clarkii was present than in places without P. clarkii , regardless of fish presence. After accounting for habitat variables and fish, crayfish presence was a negative predictor of the breeding probability for all urodeles ( Pleurodeles waltl , Salamandra salamandra , Triturus boscai and T. marmoratus ) and for two anurans ( Pelobates cultripes and Bufo bufo ). The majority of the species affected usually breed in temporary ponds without fish, but that may be colonized by the crayfish. The ongoing expansion of P. clarkii may eventually lead to a growing isolation of amphibian populations and ultimately to local extinctions and a permanent alteration of the amphibian communities in southwest Iberian Peninsula.     

The response of amphibian communities to fish and habitat features in Mediterranean permanent ponds

Generalized linear models were used to test the effect of fish, using ponds with and without fish and habitat features as covariates, on richness and abundance of amphibian species. Five fish species and six amphibian species were recorded in 60 permanent ponds located in central Italy. The choice of covariates (macrophyte cover and pond surface area) was made after studying the correlations. The richness of amphibian species was not significantly affected by fish presence or macrophyte cover, in line with previous studies, since almost all the fish species were non-predatory. However, abundance of urodeles (newts) was negatively affected by fish and positively affected by macrophyte cover. Although fish may strongly influence the abundance and composition of amphibian communities, the results indicate that the cover of aquatic macrophytes may increase the available habitat for amphibians and therefore their abundance. Anuran species preferred ponds where fish were present, since both groups preferred larger ponds. Concordance between fish and amphibian species composition was not found by the Mantel and Partial Mantel tests. This indicates that the fish assemblages do not predict which amphibian species occur in the pond.     

Presence of the Chytrid Fungus Batrachochytrium dendrobatidis in Populations of the Critically Endangered Frog Mannophryne olmonae in Tobago, West Indies

The emerging infectious disease chytridiomycosis is prevalent in Central and South America, and has caused catastrophic declines of amphibian populations in the Neotropics. The responsible organism, Batrachochytrium dendrobatidis, has been recorded on three West Indian islands, but the whole of the Caribbean region is predicted to offer a suitable environment for the disease. Monitoring the spread of chytridiomycosis is thus a priority in this region, which has exceptionally high levels of amphibian endemism. PCR analysis of 124 amphibian skin swabs in Tobago (Republic of Trinidad and Tobago) demonstrated the presence of B. dendrobatidis in three widely separated populations of the frog Mannophryne olmonae, which is listed as Critically Endangered on the basis of recent population declines. Chytridiomycosis is presently endemic in this species, with a prevalence of about 20% and no associated clinical disease. Increased susceptibility to chytridiomycosis from climate change is unlikely in amphibian populations in Tobago, as this island does not have high montane environments, but remains a possibility in the sister island of Trinidad. Preventing the spread of chytridiomycosis within and between these and other Caribbean islands should be a major goal of practical conservation measures for amphibians in the region.     

Comparison of the hydrological characteristics of three small experimental holm oak forested catchments in NE Spain in relation to larger areas

Precipitation and streamflow have been measured in three small (0.04–0.52 km²) experimental catchments covered by dense holm oak (Quercus ilex L.) forests. Two of them are in the Prades mountains and one in the Montseny mountains (NE Spain). Here we test the hydrological representativeness of these catchments against the streamflow record at two nearby larger (34–60 km²) catchments, one from each massif. Comparisons of (i) annual streamflow, (ii) seasonal distribution of streamflow, and (iii) flow duration curves were made. At Prades, for the period of common record, mean annual precipitation was about 580 mm, and mean annual streamflow 44–81 mm at the two experimental catchments and 102 mm at the larger one. Most streamflow occurred during winter and spring in the three catchments. At Montseny, rainfall was higher, and mean annual streamflow was 495 mm in the experimental catchment, and 760 mm in the larger catchment, though these data were obtained in different periods in each catchment. Streamflow was roughly equal in autumn, winter and spring. At both sites flow duration curves were fairly similar in the small experimental catchments and the larger catchments. The higher streamflow at Montseny is reflected in its flow duration curves being well above those at Prades. The experimental catchments at Prades are thus fairly representative of the larger nearby catchment for the investigated hydrological characteristics. At Montseny, hydrological differences between the experimental catchment and the larger catchment are probably due to the higher mean altitude of the latter and to the non-overlapping periods of their streamflow records.     

Amphibian diversity increases in an heterogeneous agricultural landscape

As a group amphibians are the vertebrates most affected by anthropic activity, particularly by agriculture. The rapid advance of the agricultural frontier makes it important to identify the role of agroecosystems as habitat supply for amphibians. We analyzed the differences in amphibian assemblages and populations between habitats with different plant covers and different degrees of human intervention in northwestern Argentina. For three years we conducted 114 high frequency trap samplings to quantify abundance, specific composition and species richness of amphibian assemblages in three habitat types (lemon and sugarcane crops and secondary forest) of a piedmont agroecosystem of Tucumán province. Crops hosted more species and individuals than secondary forests, but the specific composition of forest was different from that of crops suggesting that they could be complementary. Although the assemblage abundance of every observation responded to climate, the strong effect of sampling year was not related to climatic factors suggesting that there might be long term fluctuations that were not analyzed. We also found that responses to agricultural practices were species specific, so no generalizations about these practices should be done. Our study shows that cultivated areas are not hostile environments for amphibians since they can lodge huge amphibian populations and that the occurrence of disturbances associated to agricultural practices is not necessarily reflected in a decrease in the abundance and richness of amphibians in the short term. However, our results also show that forested lands are necessary to lodge some specialist species which are very rare in the croplands. This suggests that environmental heterogeneity generated by the combination of natural and cultivated patches can increase biodiversity at landscape scale because this allows the coexistence of species related to either kind of environment.     

Populations of a susceptible amphibian species can grow despite the presence of a pathogenic chytrid fungus

Disease can be an important driver of host population dynamics and epizootics can cause severe host population declines. Batrachochytrium dendrobatidis (Bd), the pathogen causing amphibian chytridiomycosis, may occur epizootically or enzootically and can harm amphibian populations in many ways. While effects of Bd epizootics are well documented, the effects of enzootic Bd have rarely been described. We used a state-space model that accounts for observation error to test whether population trends of a species highly susceptible to Bd, the midwife toad Alytes obstetricans, are negatively affected by the enzootic presence of the pathogen. Unexpectedly, Bd had no negative effect on population growth rates from 2002-2008. This suggests that negative effects of disease on individuals do not necessarily translate into negative effects at the population level. Populations of amphibian species that are susceptible to the emerging disease chytridiomycosis can persist despite the enzootic presence of the pathogen under current environmental conditions.     

Habitat associations of pool-breeding amphibians in eastern Massachusetts,USA

There is a pressing need to develop a sound conservation strategy for pool-breeding amphibians, which includes gaining a better understanding of the habitat and landscape-scale characteristics associated with populations. To investigate relationships between amphibian species richness and characteristics of breeding pools and surrounding landscapes, we surveyed 85 pools in eastern Massachusetts (USA) in 1996 and 1997. A total of 11 species was detected, with most pools having 2–5 species. Pools were typically small, 77.6% were <0.2 ha, but most pools (72%) had hydroperiods that persisted at least into August in most years. Based on linear regression analyses, species richness was positively associated with three within-pool variables (pool surface area, hydroperiod, and the amount of emergent vegetation), and a landscape-level variable (presence of another breeding pool within 1 km), while one within-pool variable (tree canopy cover) exhibited a significant negative association with species richness. These within-pool habitat variables and connectivity to other breeding pools are important characteristics to consider when attempting to identify breeding sites that could provide core habitat in conservation reserves designed for the conservation of pool-breeding amphibian species richness. Conservation reserves for pool-breeding amphibian populations should include pool complexes functioning as habitat for metapopulations. Core pool habitats within such reserves should be large (0.5–1.0 ha), with seasonal hydroperiods that persist into August or that dry in some years, and with sufficient emergent vegetation to provide diverse microhabitats and refugia.     

Size-structured effects of common carp on reproduction of pond-breeding amphibians

The role of fish in driving amphibian communities has been widely recognized. However, little is known about size-structured interactions between amphibian and fish populations. This study compared the taxonomic occurrence and densities of larval amphibians between unstocked ponds and ponds stocked with different age cohorts of common carp Cyprinus carpio differing in average body size. The average total densities of early and late breeding anurans known to be vulnerable to fish were by 1–2 orders of magnitude greater in the presence of young-of-the-year carp than that of older cohorts. The probabilities of occurrence of the most common taxa did not differ between ponds stocked with young-of-the-year fish and ponds free of carp, but were significantly larger in those ponds than in ponds stocked with large-size cohorts. No significant differences between pond categories were found for densities of unpalatable Bufo bufo larvae. In aquatic systems harbouring size-structured fish populations, a fish age/size gradient may explain differential habitat suitability for breeding amphibians better than the fish presence/absence dichotomy. When dominated by young cohorts incapable of predation or of adverse habitat alteration, fish-abundant waters are suitable for amphibian reproduction. Conversely, even a ‘non-predatory’ fish, after attaining large body size, may exert a detrimental impact on amphibian breeding success. These findings may be particularly important for amphibian conservation at pond fisheries characterized by spatial separation of age/size distributed stocks.     

Lower genetic diversity and hatchability in amphibian populations isolated by urbanization

Many amphibian populations worldwide have declined rapidly and been threatened with extinction in the past few decades because of human impacts on the environment. It is well known that urbanization reduces the genetic diversity of isolated populations. The concept that a reduction in genetic diversity leads to lower reproductive fitness has been predominantly supported by studies involving laboratory organisms, but has rarely been tested in wild populations. Here we examined whether genetic diversity affected hatchability, a population parameter related to reproductive success, in the populations of two pond-breeding amphibian species (Hynobius tokyoensis and Rana ornativentris) affected by urbanization. We surveyed 81 populations in the southwestern Kanto region of Japan. Mean hatchability of the populations was determined and genetic diversities were estimated via mitochondrial or microsatellite DNA analyses. Firstly, Random Forests (an ensemble machine learning method) models were applied to clarify the effects of environmental factors on both hatchability and genetic diversity. Subsequently, the relationships among environmental factors, genetic diversity, and mean hatchability were evaluated using path analysis. Mean hatchability was significantly affected by both urbanization and genetic diversity. This result shows that loss of genetic diversity may decrease a population’s reproductive success in the field.     

Canopy closure and amphibian diversity in forested wetlands

The crowns of trees and shrubs often overtop temporary wetlands in forested regions. By shading pond basins, canopy can dramatically change the conditions experienced by residents such as amphibians. In this study, we estimated the presence of 8 amphibian species across 17 temporary wetlands at the Yale–Myers Forest in northeastern Connecticut, USA. In addition, we quantified the light environment using a grid of hemispherical canopy photographs to calculate Global Site Factor (GSF) within each wetland. Amphibian richness was low in most wetlands, and most wetlands were relatively shaded. Amphibian richness increased in lighter wetlands. This result was not confounded by relationships with wetland size. Most amphibian species tended to be absent from heavily shaded wetlands (‘open canopy specialists’). However, three species were often found in the shadiest wetlands (‘canopy generalists’). Field transplant experiments using one canopy generalist and one open canopy specialist showed that development of the generalist was less affected by wetland light levels compared with performance of the specialist. These findings suggest that canopy may be an important determinant of amphibian diversity patterns across wetlands. Further, conservation strategies dependent on universally applied, inviolate shoreline vegetation buffers may inadvertently contribute to species loss. Because species differ in their sensitivity to changes in canopy, these losses may be predictable.     

Analysis of three amphibian populations with quarter-century long time-series.

Amphibians are in decline in many parts of the world. Long tme-series of amphibian populations are necessary to distinguish declines from the often strong fluctuations observed in natural populations. Time-series may also help to understand the causes of these declines. We analysed 23-28-year long time-series of the frog Rana temporaria. Only one of the three studied populations showed a negative trend which was probably caused by the introduction of fish. Two populations appeared to be density regulated. Rainfall had no obvious effect on the population fluctuations. Whereas long-term studies of amphibian populations are valuable to document population declines, most are too short to reveal those factors that govern population dynamics or cause amphibian populations to decline.     

Genetic evidence for a high diversity and wide distribution of endemic strains of the pathogenic chytrid fungus Batrachochytrium dendrobatidis in wild Asian amphibians

Population declines and extinctions of amphibians have been attributed to the chytrid fungus Batrachochytrium dendrobatidis (Bd), especially one globally emerging recombinant lineage (‘Bd‐GPL’). We used PCR assays that target the ribosomal internal transcribed spacer region (ITS) of Bd to determine the prevalence and genetic diversity of Bd in South Korea, where Bd is widely distributed but is not known to cause morbidity or mortality in wild populations. We isolated Korean Bd strains from native amphibians with low infection loads and compared them to known worldwide Bd strains using 19 polymorphic SNP and microsatellite loci. Bd prevalence ranged between 12.5 and 48.0%, in 11 of 17 native Korean species, and 24.7% in the introduced bullfrog Lithobates catesbeianus. Based on ITS sequence variation, 47 of the 50 identified Korean haplotypes formed a group closely associated with a native Brazilian Bd lineage, separated from the Bd‐GPL lineage. However, multilocus genotyping of three Korean Bd isolates revealed strong divergence from both Bd‐GPL and the native Brazilian Bd lineages. Thus, the ITS region resolves genotypes that diverge from Bd‐GPL but otherwise generates ambiguous phylogenies. Our results point to the presence of highly diversified endemic strains of Bd across Asian amphibian species. The rarity of Bd‐GPL‐associated haplotypes suggests that either this lineage was introduced into Korea only recently or Bd‐GPL has been outcompeted by native Bd strains. Our results highlight the need to consider possible complex interactions among native Bd lineages, Bd‐GPL and their associated amphibian hosts when assessing the spread and impact of Bd‐GPL on worldwide amphibian populations.     

Predator-induced delayed maturity in bluegill sunfish (Lepomis macrochirus): variation among populations

Previous studies suggested that differences in age at maturity among populations of bluegill sunfish (Lepomis macrochirus) were not genetically based, but rather were a phenotypic response to the presence of predators. I conducted two experiments to determine if the presence of largemouth bass affected age at maturity in bluegill sunfish. Bluegills from three populations were tested to see if the response to the threat of predation varied among source populations. Juvenile bluegills were maintained in the presence of predators or in controls with no contact with predators. Refuge use and growth were monitored during the experiments and reproductive activity was evaluated when bluegills reached age 1. Bluegills from one population exhibited delayed maturity in the presence of predators. Individuals from the other two populations showed no significant differences between predator and control treatments. The population that responded to the presence of predators had a history of high predation levels over the past 30–40 years. The other populations had a history of low levels of predation. This study suggests that presence of predators can induce phenotypic shifts in age at maturity of bluegills, but that the magnitude of response varies among populations in a manner consistent with historical patterns of coexistence. Received: 7 August 1996 / Accepted: 8 August 1997     

Dispersal and the metapopulation paradigm in amphibian ecology and conservation: are all amphibian populations metapopulations?

Amphibians are frequently characterized as having limited dispersal abilities, strong site fidelity and spatially disjunct breeding habitat. As such, pond‐breeding species are often alleged to form metapopulations. Amphibian species worldwide appear to be suffering population level declines caused, at least in part, by the degradation and fragmentation of habitat and the intervening areas between habitat patches. If the simplification of amphibians occupying metapopulations is accurate, then a regionally based conservation strategy, informed by metapopulation theory, is a powerful tool to estimate the isolation and extinction risk of ponds or populations. However, to date no attempt to assess the class‐wide generalization of amphibian populations as metapopulations has been made. We reviewed the literature on amphibians as metapopulations (53 journal articles or theses) and amphibian dispersal (166 journal articles or theses for 53 anuran species and 37 salamander species) to evaluate whether the conditions for metapopulation structure had been tested, whether pond isolation was based only on the assumption of limited dispersal, and whether amphibian dispersal was uniformly limited. We found that in the majority of cases (74%) the assumptions of the metapopulation paradigm were not tested. Breeding patch isolation via limited dispersal and/or strong site fidelity was the most frequently implicated or tested metapopulation condition, however we found strong evidence that amphibian dispersal is not as uniformly limited as is often thought. The frequency distribution of maximum movements for anurans and salamanders was well described by an inverse power law. This relationship predicts that distances beneath 11–13 and 8–9 km, respectively, are in a range that they may receive one emigrating individual. Populations isolated by distances approaching this range are perhaps more likely to exhibit metapopulation structure than less isolated populations. Those studies that covered larger areas also tended to report longer maximum movement distances – a pattern with implications for the design of mark‐recapture studies. Caution should be exercised in the application of the metapopulation approach to amphibian population conservation. Some amphibian populations are structured as metapopulations – but not all.     

Treatment of chytridiomycosis with reduced-dose itraconazole

Effective treatment methods to eliminate infection with Batrachochytrium dendrobatidis (Bd) are required for development of sustainable captive survival assurance populations of amphibians and to reduce the risk of introducing Bd to new locations as part of amphibian trade or reintroduction programs. Treatment with itraconazole baths at 100 mg l-1 is commonly used in captive amphibians, but side effects are observed in some amphibian species and life stages. Naturally occurring outbreaks of chytridiomycosis in Wyoming toads Anaxyrus baxteri and White's tree frogs Litoria caerulea were treated with lower-dose itraconazole baths (e.g. 50 mg l-1 for White's tree frogs) and followed post-treatment with serial Taqman PCR testing to confirm elimination of Bd infection. Post-treatment PCR tests were consistently negative for the presence of Bd and treatment was deemed successful. Although this was not a controlled clinical trial, results suggest that lower doses of itraconazole may be effective for treatment of chytridiomycosis with resulting cost savings to amphibian conservation programs and a potential for a reduction in dose-related side effects from itraconazole treatment. Prospective clinical trials of alternative itraconazole treatment protocols are encouraged.     

Can a large metropolis sustain complex herpetofauna communities? An analysis of the suitability of green space fragments in Rome

Urban areas are primary causes of species' range fragmentation and reduction. However, relatively few studies have attempted to describe the habitat variables influencing the diversity and conservation of amphibians and reptiles, particularly in Mediterranean Europe and in large metropolitan areas. We explored this broad conservation ecology problem by studying the richness and diversity patterns in relation to a suite of six independent habitat variables in Rome, one of the most ancient cities of the world. We considered all the green remnant areas ( n =62) of Rome, ranging 1 to >1000 ha in size, which are interspersed within a sea of urbanized matrix. A total of 10 amphibian and 15 reptile species were studied. Their presence/absence patterns were assessed and the effects of the various habitat variables on each species were predicted by a logistic regression model. A total of 1261 presence records (404 amphibians and 857 reptiles) were analysed. Fragment size and wood size within each fragment did correlate significantly with the species richness of both amphibians and reptiles, and there was a clear threshold effect after 50 ha of wooded surface. The presence of water bodies positively affected the species distribution. One amphibian and three reptiles inhabited exclusively fragments >50 ha. The distance from the centre did not affect fragment species richness. The presence of most species of both amphibians and reptiles was positively influenced by the irregular versus circular shape of the wooded area. The legal protection of a given area did not influence the observed patterns but the total number of sheltered species. Overall, our study suggests that, in order to maintain the current diversity and population viability, it is necessary, in addition to water bodies' maintenance, to (1) preserve the wooded landscapes over 50 ha; (2) promote irregularly shaped increases in the wood surface; (3) maintain ecotonal boundaries.     

Intersex Frogs Concentrated in Suburban and Urban Landscapes

The occurrence of intersex characteristics in amphibians has been linked to pesticide exposure in the laboratory and proximity to agricultural activity within natural populations. But, overall, the ecology of amphibian intersex is poorly known and, specifically, its occurrence in many landscape types and regions is unstudied. We offer the first analysis of the frequency of amphibian intersex across a range of land covers representing the major landscape types within a region. We used remotely sensed information to characterize land cover surrounding 4774 potential sampling locations within the Connecticut River Valley. From among these, we selected 24 ponds to collect postmetamorphic green frogs (Rana clamitans) from four land cover types: undeveloped, agricultural, suburban, and urban. Collected males were preserved and, then, prepared gonadal tissue samples were screened for the presence of testicular oocytes. A total of 233 animals was examined. Thirteen percent of all male green frogs had gonads containing testicular oocytes. Sexual abnormalities were not randomly distributed among sites or landscape types. Suburban landscapes had the highest frequency of abnormalities (21%), and both suburban and urban land covers were positively associated with the presence of abnormalities within a population. There was no evidence of a positive association with agricultural land cover. Examination of amphibian intersex across a range of contexts reveals that developed landscapes may be hotspots for abnormal sexual development. This new finding suggests that other mechanisms, not previously considered, could contribute to intersex in natural amphibian populations.     

Habitat-mediated impact of alien mink predation on common frog densities in the outer archipelago of the Baltic Sea

Alien predators have been recognised as one possible cause for amphibian declines around the world, but little is known of habitat-mediated predation impacts especially on adult amphibians. We studied common frog Rana temporaria under American mink Mustela vison predation in the outer archipelago of the Baltic Sea, south-western Finland. Using egg batches as an index of breeding frog female numbers we compared frog numbers and densities between a large, long-term mink-removal area and a comparable control area. Frog numbers in the removal area were at least 2.7-fold higher than those in the control area. In the presence of mink, frog densities increased with the amount of vegetation cover on the islands, indicating that mink predation affected frog densities especially on less-vegetated islands. An opposite trend appeared to be true for frogs in the mink-removal area, where other predators like snakes could induce a decline of frog densities on more vegetated islands. Shrub or grass vegetation seems to provide frogs shelter against alien mink predation. Our result highlights the importance of landscape-level habitat management as a conservation tool for amphibian populations.