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.     

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.     

Detrimental effects of peat mining on amphibian abundance and species richness in bogs

Peat bogs of eastern North America are increasingly being mined for peat, which results in the extensive draining of these systems. First, I quantified the effects of peat mining on amphibian species richness and abundance in bog remnants adjacent to mining activity and unmined bogs. I then investigated amphibian occurrence patterns in bog remnants relative to the distance to the mined edge, after accounting for microhabitat and distance to landscape elements. The response of amphibians to peat mining varied across taxa. For instance, amphibian species richness, total amphibian captures (all spp. combined), as well as those of green frogs (Rana clamitans) were greater in unmined bogs than in bog remnants. Wood frogs (Rana sylvatica) were most abundant in unmined bogs in areas far from bog ponds, as well as in unmined bogs in areas near shrub patches. After accounting for microhabitat and landscape variables, American toads (Bufo americanus) in bog remnants were most abundant close to mined edges, whereas the other species did not respond to the proximity of mined edges. Peat mining negatively impacts amphibians occurring in bog remnants. The response of amphibians to peat mining is similar to the one of birds and plants, but differs with small mammals. Maintaining a complex mosaic of bog ponds, shrub and forest patches might help mitigate peat mining effects on amphibian patterns of abundance.     

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.     

No evidence of general decline in an amphibian community of Southern France

We investigated long term changes in an amphibian community in the Languedoc area of Southern France by comparing results of a survey of 56 ponds made in the early 1970s with results of a survey of the same localities in 2001. Based on the frequency of new occurrences and disappearances of species in the sample ponds, there is no sign of a general decline of amphibians in this area. Most species showed non-significant variation in frequency of occurrence or had increased. “Green frogs” (Rana perezi/ridibunda/kl. grafi) showed clear signs of decline, but this is a local phenomenon rather than a decline at the regional scale. Pelodytes punctatus probably declined also, possibly due to its sensitivity to fish introduction. The species richness in the ponds has also remained stable except in ponds where fish have been introduced. The only adverse impact of human activities that we could detect was thus the introduction of fish, which had a significantly adverse effect on amphibian species richness.     

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.     

Sex and seasonal differences in the spatial terrestrial distribution of gray treefrog (Hyla versicolor) populations

Fragmentation of terrestrial landscapes has exacerbated the need to understand the spatial requirements of organisms, especially those that undergo seasonal migrations, such as pond-breeding amphibians. Pond-breeding amphibians spend much of their lives in terrestrial habitat at some distance from aquatic breeding sites. The terrestrial habitat required by a particular individual encompasses the area adjacent to a breeding pond that is used for activities such as foraging or overwintering, as well as the expanses of habitat through which it moves. To elucidate amphibian spatial habitat requirements, we monitored gray treefrog (Hyla versicolor) movements through terrestrial habitat adjacent to breeding ponds using radiotelemetry and mark–recapture along transects of artificial refugia. Results indicate that gray treefrogs frequently make migrations between foraging grounds, overwintering sites, and breeding ponds of 200 m or more, with a maximum-recorded movement distance of 330 m. Additionally, females travel farther than males for reasons independent of body size, and the distribution of males during the breeding season is biased towards breeding ponds relative to the non-breeding season. The data illustrate the importance of habitat directly adjacent to breeding sites for males, and indicate that habitat loss resulting in small patches may have a greater negative impact on females than males, and consequently may have disproportionately large effects on population persistence. These results emphasize that effective management of amphibian breeding locations via protection of both aquatic and adjacent upland terrestrial resources likely requires detailed information regarding variation in movements between the sexes and among seasonal activity periods.     

Distribution of the Mallorcan midwife toad (Alytes muletensis) in relation to landscape topography and introduced predators

The endemic midwife toad of Mallorca (Alytes muletensis) is restricted to a small number of breeding populations in the mountainous northwest of the island. The decline of the species has been attributed to the impacts of introduced species such as the viperine snake (Natrix maura) and green frog (Rana perezi), and toads may be surviving only in areas that are suboptimal for these predators. The influence of landscape features (elevation, aspect and maximum slope) on the distribution of toads and associated predators was therefore investigated using GIS. The presence of toads was positively associated with steep slopes. At sites where they occurred with toads, the distribution of predators was negatively associated with elevation. Reproductive success within toad populations was strongly associated with the number of pools at each site, while reproductive success within individual pools was positively associated with elevation. These findings may be used to optimise the design and location of future reintroduction sites.     

Mass mortality of native anuran tadpoles in tropical Australia due to the invasive cane toad (Bufo marinus)

Specific pathways of the ecological impact of invasive species remain poorly known. Although the spread of toxic cane toads (Bufo marinus) through tropical Australia is widely believed to have caused extensive mortality of native reptiles and mammals, effects of toad ingestion on native anurans have been virtually ignored. Our studies on the Adelaide River floodplain show that the most numerous vertebrate victims of toad invasion are native tadpoles that die when they attempt to consume toad eggs. We documented 11 episodes of mass mortality, totalling >1300 tadpoles of 10 species, in five waterbodies within a single wet-season shortly after the toads invaded. A causal link between toad breeding and tadpole mortality is supported by observations that: (1) in at least 9 of the 11 waterbodies involved, toads bred immediately prior to mortality events; (2) water quality was indistinguishable from that of control ponds, and tadpoles placed in that water remained healthy; (3) dead tadpoles showed no sign of disease; and (4) laboratory trials showed rapid, 100% mortality in native tadpoles exposed to freshly-laid toad eggs. Despite these high mortality rates, toad invasion does not appear to threaten the viability of anuran populations because frogs often breed in ponds not used by toads, and because density-dependent growth and survival within tadpole communities mean that additional mortality may not reduce the total effective recruitment of metamorph frogs from a waterbody.     

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.     

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.     

A survey of natterjack toad Bufo calamita distribution and breeding success in the north merseyside sand-dune system,England

In spring 1978, adult natterjack toads Bufo calamita assembled at 101 out of 132 sites containing water in the north Merseyside sand-dunes. A total of 2001 spawn strings was counted in 95 sites, 95% of the spawn being deposited before 11 May. Metamorphosis was confirmed at 28 sites and probably occurred at a further 12 sites distributed throughout the dunes. Most spawning took place in shallow water in slacks or excavated scrapes. Deep ponds, ditches and bomb craters were little used. Unsuccessful breeding was mainly due to shallow water bodies drying up before metamorphosis could occur. Of sites where metamorphosis was confirmed, 82% were scrapes which had been excavated for conservation purposes.Water-table trends in the dune system and various methods of censusing the adult toad population are discussed.     

Abundance, body size and movement patterns of a tropical treefrog in continuous and fragmented forests in the Brazilian Amazon

Phyllomedusa tarsius is a hylid frog that breeds in ponds located in a range of habitats from continuous forests to severely disturbed matrix habitats in Central Amazon. During three reproductive seasons, we followed the movement patterns, measured body size and registered abundance and residency time of this species in five habitats: pasture, Vismia regrowth, Cecropia regrowth, 1 and 10 ha forest fragments, and continuous forest. The frog captures were strongly correlated with rainfall in all disturbed habitats, but not in continuous forest, probably because individuals respond more directly to rainfall patterns in more open habitats. Males in disturbed habitats were smaller than those found in continuous forest, perhaps as a result of differences in habitat quality. Males remained at sites for longer periods in fragments and continuous forest compared to matrix habitats. Here again, the quality and suitability of breeding sites in matrix habitats may be lower than in continuous forest resulting in the need for more movement. We found bigger subpopulations in matrix habitat ponds and a higher percentage of individuals moving among them when compared to continuous forest ponds. Constant movement of individuals among disturbed subpopulations decreases population isolation and increases interbreeding among different subpopulations. No movement between individuals from continuous forest ponds and disturbed habitats was observed. This suggests that the disturbed habitat subpopulations are not acting as sink subpopulations for continuous forest populations. The maintenance of individuals in fragments is more dependent on local conditions for reproduction and on subpopulations in the matrix habitat than on recruitment of individuals from populations in continuous forest.     

Human land use and patterns of parasitism in tropical amphibian hosts

Landscape alterations by humans can change patterns of parasite transmission. Depending on the type alteration and the life histories of parasites and hosts, parasitism may increase or decrease. To investigate whether parasitism in tropical amphibians was associated with land use change, I studied three species of amphibians, Rana vaillanti, Eleutherodactylus fitzingeri, and Smilisca puma from the Province of Heredia, Costa Rica, in the Atlantic lowlands. Frogs were collected and examined for parasites during the rainy seasons of 2001-2003 from sites in forest or clear-cut cattle pastures. The abundances of five species of parasites and parasite species richness of R. vaillanti were significantly higher in pasture habitat. A single parasite species shared by E. fitzingeri and S. puma showed a trend of increased abundance in pasture habitats. Overall, the abundances of six parasite species (three trematodes, an intestinal nematode, an encysted nematode, and a filarial nematode) were higher in pastures, while two species (trematodes) were higher in forest. This study suggests that land use activities in tropical regions affects the abundance and richness of amphibian parasites. Converting forest to pasture may impact the abundance of amphibian parasites because subsequent water quality changes (e.g. eutrophication, higher pH) often enhance habitat and resources for intermediate hosts (e.g. snails, mosquitoes) involved in parasite life cycles. Aquatic amphibians (e.g. R. vaillanti) may be particularly prone to experiencing increases in parasitism in agricultural habitats due to the positive relationship between many aquatic intermediate hosts and water quality changes associated with agricultural land use.     

Effects of increased salinity and an introduced predator on lowland amphibians in Southern China: Species identity matters

Approximately 30% of amphibian species are threatened due to a variety of factors affecting their habitats and physiology, yet contributions that interactions among factors make to population declines are not well-explored. Two factors, introduced mosquitofish (Gambusia affinis) and increased salinity, may coincide in lowland habitats used by breeding amphibians. Mosquitofish have been introduced worldwide and can be significant predators of amphibian larvae. Salinization of wetlands is an increasing problem globally due to (1) application of road deicing salts in temperate regions, (2) irrigation practices associated with intensive agriculture, and (3) saltwater intrusion due to sea-level rise. We investigated the effects of mosquitofish (G. affinis) and increasing salinity on five species of lowland wetland-breeding amphibians from southern China. We exposed anuran hatchlings to four salinity levels and two fish treatments and all combinations of the two factors in a series of experiments. Four of the species were susceptible to predation by mosquitofish, two were sensitive to increased salinity at concentrations >6% seawater, and one was tolerant of both increased salinity and mosquitofish. We found no interaction between the predator and increased salinity. Salinization and mosquitofish represent significant threats to lowland amphibians in this region and, coupled with the ongoing loss and degradation of lowland wetlands, portend a bleak future for lowland amphibian populations in the region.     

Acid deposition from the Russian Kola Peninsula: Are sensitive fish populations in north-eastern Finnish Lapland affected?

Sulphur emissions from Russian Kola Peninsula smelters are known to cause surface water acidification in the border areas between Norway and Russia. The sulphur deposition is also high in the eastern part of Finnish Lapland. In 1990, a monitoring programme was started to survey the effects of acid deposition on sensitive fish populations in north-eastern Finnish Lapland. Altogether 103 sites in three areas were electrofished and autumn water samples were taken. Besides the brown trout (Salmo trutta), special attention was paid to the occurrence of minnow (Phoxinus phoxinus) since it is a common species in small waters and is highly sensitive to acidification. During the first three years of monitoring no signs of acidification were recorded. The alkalinity values of brooks generally exceeded 0.1 mmol/1. Brown trout, minnow and burbot (Lota lota) were caught frequently in the study sites. Later the study was focused on the uninhabited Vätsäri area which is receiving the highest sulphur deposition in Finnish Lapland. The alkalinity values of the sampled brooks were in most cases below 0.05 mmol/1, indicating a decreased buffer capacity. However, the electrofishing of the brooks showed no acid-induced damage. The lowest alkalinity values were detected from a group of small upland ponds. In four such ponds the alkalinity was zero or negative. No minnows were caught from these four ponds apart from one, where the minnows were exceptionally large. The results show that the waters near the eastern border of northern Finnish Lapland are threatened by acidification. No damage to fish populations subject to fishing was observed. The absence of minnows in some small waters is possibly the first sign of acid-induced fish population damage.     

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.     

Recent, substantial, and unexplained declines of the common toad Bufo bufo in lowland England

A decline in numbers of a large common toad (Bufo bufo) population in south-east England during the 1990s, together with anecdotal reports of similar trends in other toad populations, prompted a nation-wide survey of this species. The survey also included the common frog (Rana temporaria) as a control for which there was no comparable evidence of recent decline. A questionnaire requesting information on the fate of toad and frog populations in the last 15 years of the 20th century was distributed to professional and amateur herpetologists during autumn 2001. Ninety-five respondents provided data on a total of 277 sites, including 232 frog and 202 toad populations in England, Scotland and Wales. More than 80% of the reported sites were rural for both species. Rural frog populations were generally doing well in most parts of Britain, with almost as many increasing as decreasing populations and a high proportion of stable populations. Rural toads also showed no overall trend across the country as a whole. However, regional analysis indicated that although toads were faring at least as well as frogs in the north and west of Britain, they were declining seriously in central and eastern/south-eastern areas. In these regions 50% or more of toad populations have experienced recent declines, whereas frogs have fared as well as they have elsewhere. Toads breeding alone have fared significantly worse than toads breeding at sites where frogs were also present. Reasons for apparently toad-specific declines in lowland England remain unknown.     

Distribution,abundance and conservation of the bald ibis Geronticus calvus in Southern Africa

Updated information is provided on the breeding distribution and abundance of the bald ibis Geronticus calvus in southern Africa. Results of long-term, comprehensive surveys of the species in the Transvaal and Orange Free State are presented, and combined with information on the bird's breeding abundance in Natal to estimate its total breeding population in South Africa. The bald ibis's status in Swaziland and Lesotho is also examined. The South African breeding population is estimated at approximately 3440 birds, while its total breeding population may exceed 4600 birds. Human disturbances at its breeding sites, and degradation of its grassland biotope, pose the greatest threats to the bald ibis. Recommendations for improving the conservation of the species are discussed.