Effects of Experimental Canopy Manipulation on Amphibian Egg Deposition

ABSTRACT Although effects of forest management on amphibians are relatively well studied, few studies have examined how these practices affect egg deposition by adults, which can impact population recruitment. We quantified the effects of 4 canopy tree-retention treatments on amphibian oviposition patterns in clusters of 60-L aquatic mesocosms located in each treatment. We also related aquatic and terrestrial biophysical parameters in treatment plots to oviposition patterns. Cope's gray treefrogs (Hyla chrysoscelis) deposited more egg masses in clear-cut and 25–50% tree-retention treatments than in controls. In contrast, mountain chorus frogs (Pseudacris brachyp***hona) deposited more egg masses in unharvested control and 75% retention treatments than in clear-cut or 25–50% retention treatments. Spotted salamanders (Ambystoma maculatum) only deposited eggs in 75% retention treatments and controls. The number of egg masses deposited by mountain chorus frogs was positively related to canopy cover and negatively related to water temperature, pH, and dissolved oxygen, whereas we noted the opposite relationships for Cope's gray treefrogs. We did not detect a relationship between the number of egg masses deposited by any species and the distance of mesocosms to either the nearest mature closed-canopy forest or to the nearest natural amphibian breeding pool. The impacts of the silvicultural treatments we studied were species-specific and depended on the amount of trees removed. In areas where protection of spotted salamander and mountain chorus frog breeding habitat is a priority, we recommend harvests retain at least 75% of the canopy. Our results also suggest that retention of 25–50% of canopy trees surrounding amphibian breeding pools has little conservation benefit.     

Flexible oviposition strategies in túngara frogs and their implications for tadpole spatial distributions

Because oviposition site selection is often studied in a single ecological context, little is known about flexibility in oviposition strategies. We studied the oviposition site selection strategies of túngara frogs ( Physalaemus pustulosus ) with respect to conspecific eggs and larvae in two different ecological contexts-arrays of artificial ponds and natural stream pools. We fit maximum likelihood models to sequential pond use data for both systems and found that strategies for oviposition differed greatly between the two systems. Within arrays of artificial ponds, túngara frogs avoided both conspecific egg masses and conspecific larvae. However, within a set of natural stream pools, túngara frogs consistently oviposited in the same pools from night to night, while many nearby ponds went unused. These variable strategies produced very different distributions of tadpoles among ponds, such that tadpoles tended to be evenly spread among artificial ponds, but highly clumped within stream pools. The difference between the two strategies appears to be an overriding effect of habitat quality. Surface area and the amount of leaf litter were all significant predictors of whether stream pools were used. Additionally, an experiment in which egg masses were either transplanted to, or removed from, selected stream pools provided no evidence that frogs in these pools used egg masses per se as a cue for oviposition. These results illustrate that oviposition strategies may be very flexible, allowing species to adjust strategies to different ecological contexts. Furthermore, they demonstrate that variation in oviposition strategies may have strong effects on egg and larval spatial distributions in patchy environments.     

Double-observer approach to estimating egg mass abundance of pool-breeding amphibians

Interest in seasonally flooded pools, and the status of associated amphibian populations, has initiated programs in the northeastern United States to document and monitor these habitats. Counting egg masses is an effective way to determine the population size of pool-breeding amphibians, such as wood frogs (Rana sylvatica) and spotted salamanders (Ambystoma maculatum). However, bias is associated with counts if egg masses are missed. Counts unadjusted for the proportion missed (i.e., without adjustment for detection probability) could lead to false assessments of population trends. We used a dependent double-observer method in 2002–2003 to estimate numbers of wood frog and spotted salamander egg masses at seasonal forest pools in 13 National Wildlife Refuges, 1 National Park, 1 National Seashore, and 1 State Park in the northeastern United States. We calculated detection probabilities for egg masses and examined whether detection probabilities varied by species, observers, pools, and in relation to pool characteristics (pool area, pool maximum depth, within-pool vegetation). For the 2 years, model selection indicated that no consistent set of variables explained the variation in data sets from individual Refuges and Parks. Because our results indicated that egg mass detection probabilities vary spatially and temporally, we conclude that it is essential to use estimation procedures, such as double-observer methods with egg mass surveys, to determine population sizes and trends of these species.     

Up-regulation of glutathione metabolism and changes in redox status involved in adaptation of reed (Phragmites communis) ecotypes to drought-prone and saline habitats

The glutathione (GSH) metabolic characteristics and redox balance in three ecotypes of reed (Phragmites communis), swamp reed (SR), dune reed (DR), and heavy salt meadow reed (HSMR), from different habitats in desert regions of northwest China were investigated. The DR possessed the highest rate of GSH biosynthesis and metabolism with the lowest levels of total and reduced GSH and its biosynthetic precursors, gamma-glutamylcysteine (gamma-EC) and cysteine (Cys), of the three reed ecotypes. This suggests that a higher rate of GSH biosynthesis and metabolism, but not GSH accumulation, might be involved in the adaptation of this terrestrial reed ecotype to its dry habitat. The HSMR shared this profile although it exhibited the highest reduced thiol levels of the three ecotypes. Two key enzymes in the Calvin-cycle possessing exposed sulfhydryl groups, NADP(+)-dependent glyceraldehydes-3-phosphate dehydrogenase (G3PD) and fructose-1,6-bisphosphatase (FBPase), and other two key enzymes in the pentose-phosphate pathway (PPP), glucose-6-phosphate dehydrogenase (G6PDH) and 6-phosphogluconate dehydrogenase (6-PGD), had very similar activities in the three reed ecotypes. Compared to the SR, the DR and HSMR had higher ratios of NADPH/NADP+ and NADH/NAD+, indicating that a more reduced redox status in the plant cells might be involved in the survival and adaptation of the two terrestrial reed ecotypes to long-term drought and salinity, respectively. These results suggest that changes of GSH metabolism and redox balance were important components of the adaptation of reed, a hydrophilic plant, to more extreme dune and saline habitats. The coordinated up-regulations of the rate of GSH biosynthesis and metabolism and reduction state of redox status of plant cells, conferred on the plant high resistance or tolerance to long-term drought and salinity.     

Amphibian‐mediated nutrient fluxes across aquatic–terrestrial boundaries of temporary wetlands

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Seasonal changes of polyamines in habitat adaptation of different ecotypes of reed plants

The leaves of four reed ecotypes (Phragmites communis Trinius) growing in the desert regions of northwest China were investigated for levels of polyamines and activity of arginine decarboxylase (ADC; EC 4.1.1.19) during the growing season of 5 months. The polyamines in the leaves of all reed ecotypes consisted of putrescine, spermidine and spermine. The polyamine levels of the leaves were lower in the swamp reed than in the terrestrial reed ecotypes. Leaf polyamine levels decreased in all ecotypes over the course of the season. Compared to the swamp reed, the terrestrial reed ecotypes maintained higher ADC activity and a predominance of spermine, resulting in a lower ratio of putrescine to spermidine and spermine. It seems that the adaptation of reed plants to drought and saline habitats may be correlated with putrescine synthesis via the ADC pathway, and with a successful conversion of putrescine to spermidine and spermine.     

Genetic variation in relation to demography of peripheral pool frog populations (Rana lessonae)

Summary Genetic variation in an isolated northern metapopulation of the pool frog (Rana lessonae) in Sweden was compared to that of Central European populations using enzyme electrophoresis and literature data. Of the 31 loci scored, two (EST-2 andIDH-2) were polymorphic while no variation occurred in seven of the eight loci which are polymorphic in Central European populations.The heterozygosity level of the Swedish pool frogs is very low compared to that of other anuran populations, but their mean proportion of fertilized eggs within egg masses (97.5%) was not lower than in more heterozygous species, and their body size-specific fecundity did not differ from that of Polish conspecifics. The low genetic variability of the Swedish pool frogs is discussed in relation to features of the local populations such as size (N), calculated effective size (N _e ) reproductive success and probable history. It is concluded that long-term strong fluctuations in population size caused by reporductive failure in cold years have contributed more to the low genetic variability than could a single founder event due to a recent introduction by man.     

The choice of oviposition site and the effects of density and oviposition timing on survivorship in<Emphasis Type="Italic"> Rana japonica</Emphasis>

In species that lack parental care and whose larvae are restricted to a given space throughout development, choice of egg-laying or reproductive sites often greatly influences fitness. Furthermore, reproductive success of individuals may be affected not only by conditions of the breeding area but also by the time when they start to breed, because intraspecific variation in the timing of breeding may result in asymmetric competition between larvae. We addressed these issues in an experimental and field study of the Japanese brown frog, Rana japonica. We tested whether the survival success of larvae decreased with an increase in the number of deposited egg masses and with a greater delay in oviposition. We found that the number of egg masses and timing of oviposition together significantly predicted larval survival per clutch. Moreover, we observed the natural oviposition of R. japonica to examine the prediction that if the density of larvae and the timing of oviposition affect survivorship of larvae, R. japonica will avoid depositing eggs in pools in which egg masses have already been deposited. We found that for small pools only, R. japonica tended to favor oviposition at unoccupied pools (those lacking egg masses and larvae) in the spawning season. For large pools, however, adults favored those already occupied by egg masses in the spawning period. The density of egg masses and tadpoles or the timing of oviposition may have less effect on spawning or the survival of tadpoles in large pools than in small pools.     

Green Frogs Show Reduced Foraging Success in Habitats Invaded by Japanese knotweed

Habitat loss is causing amphibian population declines worldwide, so there is increased attention to forces that degrade remaining habitats. Terrestrial habitats surrounding wetlands are critical foraging areas for temperate anurans. We investigated plant community changes in two old fields invaded by Japanese knotweed (Fallopia japonica) and the foraging success of Green frogs (Rana clamitans) in invaded and non-invaded portions of those fields. Within each field, vegetation data were recorded in quadrats located along two transects bisecting the invasion fronts. We placed frogs in ‘foraging buckets’ along transects and measured their change in mass over a 38 h period. There were significant changes in vegetation structure and composition associated with Japanese knotweed invasion. Diverse assemblages of native plants that covered non-invaded plots were absent from areas invaded by Japanese knotweed. There was also a significant change in vegetation architecture between invaded and non-invaded habitats. Change in frog mass declined significantly along transects, with most frogs in non-invaded plots gaining mass and no frogs in invaded plots gaining mass. Most frogs from non-invaded plots but only two from invaded plots defecated shortly after removal from foraging buckets (verification of recent feeding). We hypothesize that Japanese knotweed invasions degrade terrestrial habitat quality for frogs by indirectly reducing arthropod abundance. Nonnative plant invasions may be another factor contributing to amphibian population declines.     

An expert-based landscape permeability model for assessing the impact of agricultural management on amphibian migration

The persistence of pond-breeding amphibian populations is influenced on different spatial scales ranging from the individual breeding pond to surrounding habitat patches to landscape clusters of breeding pond populations. The connectivity among breeding ponds as well as the availability of suitable terrestrial habitats surrounding the ponds plays a major role in long-term viability of amphibian species. Besides road traffic and urban structures agricultural land activity can disrupt landscape connectivity through the use of pesticides, fertilizers and physical activity such as tillage. We developed an expert-based model to assess the impact of agricultural management measures on the migration area and terrestrial habitat availability for seven amphibian species. The model is based on a Habitat Suitability Index to identify suitable terrestrial habitats and includes a landscape permeability approach. Size of migration areas, connectivity of breeding ponds and number of reachable terrestrial habitats were modeled considering species-specific migration ranges and habitat preferences. We consider how pesticides application might lead to fragmentation and isolation of amphibian breeding pond populations. Therefore the potential disrupting impact of pesticides was simulated by inflating landscape resistance to medium and high migration cost. One amphibian species showed a decrease of migration area by 48.3% and a decrease of reachable terrestrial habitats by 41.5% at high migration costs. Three additional species showed a decrease of their migration areas between 31.5 and 35.7%. At increased migration cost, some of the investigated populations were isolated at breeding pond level or restricted to pond clusters. Our model could be used to prioritize conservation efforts for pond-breeding amphibians with adequate consideration of agricultural land use and its impact on amphibian migration.     

Patterns of reproductive‐mode evolution in Old World tree frogs (Anura,Rhacophoridae)

The Old World tree frogs (Anura: Rhacophoridae), with 387 species, display a remarkable diversity of reproductive modes – aquatic breeding, terrestrial gel nesting, terrestrial foam nesting and terrestrial direct development. The evolution of these modes has until now remained poorly studied in the context of recent phylogenies for the clade. Here, we use newly obtained DNA sequences from three nuclear and two mitochondrial gene fragments, together with previously published sequence data, to generate a well‐resolved phylogeny from which we determine major patterns of reproductive‐mode evolution. We show that basal rhacophorids have fully aquatic eggs and larvae. Bayesian ancestral‐state reconstructions suggest that terrestrial gel‐encapsulated eggs, with early stages of larval development completed within the egg outside of water, are an intermediate stage in the evolution of terrestrial direct development and foam nesting. The ancestral forms of almost all currently recognized genera (except the fully aquatic basal forms) have a high likelihood of being terrestrial gel nesters. Direct development and foam nesting each appear to have evolved at least twice within Rhacophoridae, suggesting that reproductive modes are labile and may arise multiple times independently. Evolution from a fully aquatic reproductive mode to more terrestrial modes (direct development and foam nesting) occurs through intermediate gel nesting ancestral forms. This suggests that gel nesting is not only a possible transitional state for the evolution of terrestriality, but also that it is a versatile reproductive mode that may give rise to other terrestrial reproductive modes. Evolution of foam nesting may have enabled rhacophorids to lay a larger number of eggs in more open and drier habitats, where protection from desiccation is important. Terrestrial direct development allows frogs to lay eggs independent of bodies of water, in a diversity of humid habitats, and may represent a key innovation that facilitated the evolution of nearly half of all known rhacophorid species.     

Assessing the value of secondary forest for amphibians: <Emphasis Type="Italic">Eleutherodactylus</Emphasis> frogs in a gradient of forest alteration

Secondary forests constitute a growing portion of forested areas worldwide. They might have a substantial role for the conservation of biodiversity in tropical areas, but there is little information on their potential to support forest species and the recovery of faunal communities. We studied two forest frogs (Eleutherodactylus diastema and E. fitzingeri) in an area of Costa Rica composed of a mosaic of primary forest, young secondary forest and pasture, and we compared the density of calling males in areas with different forest alteration. Autoregressive models were used to compensate for potentially undesired effects of spatial autocorrelation and pseudoreplication. Both species were most abundant in riparian, primary forest. However, E. fitzingeri was also abundant in riparian secondary forests, and its density far from the river was similar in primary and secondary forest, suggesting that river proximity can influence the recovery of secondary forest for amphibians. Conversely, the density of E. diastema was similar in secondary forest and pasture, stressing interspecific differences for recovery rate. These frogs have a keystone role in nutrient cycling and food webs, and their prompt recovery might represent an important step for the functional recovery of forests. Nevertheless, the strong interspecific differences stress the complexity of these processes.     

Functional response and area‐restricted search in a predator: seasonal exploitation of anurans by the European polecat,Mustela putorius

A study of the feeding habits and movements of 11 radiotracked polecats Mustela putorius in western France revealed that seasonal predation upon agile frogs, Rana dalmatina, was directly influenced by prey abundance and distribution. Although dietary structure showed the importance of mammalian prey (71.5%), polecats exploited nocturnal, terrestrial anurans in spring (31.6%). The periodic activity of anurans at spawning sites led both to a maximum density in spring and to a patchy distribution. The monthly variations in anuran dietary occurrences were associated with changes in frog availability. The functional response of polecats to frog density was sigmoidal shaped (type 3 response). Frog consumption rate increased more slowly than prey density but frogs were actively removed at higher density. It is therefore suggested that frog populations were moderately affected by the predator and this density dependent effect tends to stabilise anuran populations. Predation upon anurans was also correlated with a prey dispersion index as revealed by a polynomial regression. Polecats concentrated their predation on spawning congregations of the breeding adult frogs. Movements were smallest in spring and polecats changed their track length by increasing the difference between a succession of small movements and of longer journeys towards profitable sites. Changes in movements correlated with the anuran dispersion index and the response was sigmoidal (polynomial regression) revealing an area‐restricted search. This response may be regarded as an ‘aggregative response’ according to the first part of the definition of Begon et al. (1996) . Functional and area‐restricted search responses to the frog abundance and dispersion constitute an original example of predator‐prey coexistence strategies among vertebrates. I suggest that such predation could be favoured by the individualistic habits of the mustelid.     

Effects of exotic <Emphasis Type="Italic">Spartina alterniflora</Emphasis> on the habitat patch associations of breeding saltmarsh birds at Chongming Dongtan in the Yangtze River estuary,China

Smooth cordgrass (Spartina alterniflora) is one of the most invasive exotic plants of saltmarshes worldwide. To understand the effects of smooth cordgrass invasion on the habitat use and selection by breeding saltmarsh birds, we compared species number and abundance of breeding birds in native reed (Phragmites australis) and smooth cordgrass-invaded habitats (reed-cordgrass mixed habitats and cordgrass monocultures) at Chongming Dongtan in the Yangtze River estuary, China. We further examined the similarity of bird communities in different habitats and habitat selection by dominant bird species. For saltmarsh generalists, species number and abundance did not differ among the habitats. For saltmarsh specialists, species number and abundance did not differ in reed monocultures and reed-cordgrass mixed habitats but were significantly lower in cordgrass monocultures than in reed monocultures and reed-cordgrass mixed habitats. ANOSIM indicated that the difference in bird communities was larger between cordgrass monocultures and the habitats with reed than between the habitats with reed. The saltmarsh specialists preferred reed monocultures, while saltmarsh generalists avoided reed monocultures. Most species indicated no selection (neither preferred nor avoided) on reed-cordgrass mixed habitats, and no species preferred the cordgrass monocultures. The use of cordgrass monocultures by the common saltmarsh birds was negatively related to their body size. This study suggests that the spread of exotic smooth cordgrass has greatly affected the species composition and structure of local bird communities and has been especially disadvantageous to the saltmarsh specialists.     

衡阳盆地土壤动物多样性研究

对衡阳盆地紫色页岩土壤 9种生境中的土壤动物进行了调查研究 ,共获得土壤动物35 85只 ,隶属于 4门 8纲 39目 ,其中线虫类、蜱螨类为优势类群 ,占总捕量的 5 0 2 4 % ;常见类群 1 2类 ,占总捕量的 4 4 0 1 %。从土壤动物水平分布上看 ,9种生境相似程度较大 ,其中尤以芦竹林和桃树林、阔叶林和桉树林、芦竹林和桉树林之间相似程度更突出。从土壤动物垂直分布上看 ,9种生境中只有桃树林、柑桔林垂直分布较为明显 ,而各种生境中土壤动物具有明显的表聚性。芦竹林、柑桔林、桉树林、菜地、阔叶林和桃树林 6种生境土壤动物群落的多样性及均匀度呈基本一致的趋势 ,且各生境间差别不大     

Genetic and developmental studies of albino chorus frogs

Albino (amelanic) adult chorus frogs (Pseudacris triseriata) occurred with frequencies of 7 percent in 1981 and 12 percent in 1982 in breeding aggregations at a pond in the foothills of the Colorado Front Range. Laboratory matings and examination of albino egg masses suggest that the absence of melanin is due to a recessive allele. The albino phenotype displayed no deficiencies in survival of embryos, rates of embryo or larval development, or rates of growth of juvenile frogs. The absence of abnormalities in development or growth suggests that the a allele in P. triseriata has an action different from albino alleles studied previously in anurans.     

Defining core habitat of local populations of the gray treefrog (Hyla versicolor) based on choice of oviposition site

Concern over amphibian population declines and loss of terrestrial and aquatic habitat have emphasized the need to define habitat requirements for each stage in a species' life history. The realization that pond-breeding amphibians spend most of their lives in the terrestrial environment suggests the need to protect terrestrial as well as aquatic habitat. Many studies on amphibian populations have focused on emigration from breeding sites to define habitat use; however these studies do not typically elucidate terrestrial activities of adults within the breeding season. We measured colonization rates of artificial pools by gray treefrogs (Hyla versicolor) at multiple distances from natural breeding ponds. We found a non-random distribution of egg deposition among distances, with 95% of eggs deposited within 15 m of the breeding pond. Additionally, we found that the time to first colonization of artificial pools increased with respect to distance. Our results indicate that adult gray treefrogs may travel up to 200 m within a breeding season, and that multiple breeding ponds may be considered part of a single population. We suggest that a minimum core terrestrial habitat of 60 m surrounding breeding sites is appropriate for protection of local populations of gray treefrogs.     

Reedbed structure and habitat preference of reed passerines during the post-breeding period

Post-breeding habitats of nine passerine species were studied in the permanently flooded reed beds of ponds near the village Pusté úlany (SW Slovakia) in 2002. Structural features of reed beds and the abundance of all birds studied were sampled at two sites and eight study plots. The reed bed structure and abundance of four passerines differed considerably between the individual study plots. The variations in vegetation structure were also caused by winter reed burning in the SE part of study area. Stems in burnt reed beds were smaller and thinner than unburnt. Abundance of the reed warbler and to a lesser extent sedge warbler was higher in tall reed beds, while the bearded tit preferred thick reed stems. Position of reed beds along the shore — open water gradient was an important factor in terms of abundance at the study plots in the case of the reed bunting. This species showed a strong preference for the reed bed edge with open water. Horizontal distribution of other species seems to be random, however, the birds with the exception of Savi’s warbler preferred the upper stratum of reed bed in vertical distribution. Our field data corroborate with those found in experimental studies concerning with ecomorphological characteristics of birds. Some inconsistencies, however, could also be caused by other factors (e.g. foraging preferences). Comparison of breeding and post-breeding habitat requirements of the studied bird species confirmed a more or less random distribution of the majority of species in the post-breeding period. Some variation was found in the cases of only the reed warbler and bearded tit.     

Reciprocal subsidies in ponds: does leaf input increase frog biomass export?

Reciprocal subsidies occur when ecosystems are paired, both importing and exporting resources to each other. The input of subsidies increases reciprocal subsidy export, but it is unclear how this changes with other important factors, such as ambient resources. We provide a conceptual framework for reciprocal subsidies and empirical data testing this framework using a pond–forest system in Missouri, USA. Our experiment used in situ pond mesocosms and three species of anurans: wood frogs, American toads, and southern leopard frogs. We predicted that increases in ambient resources (primary productivity) and detrital subsidy input (deciduous tree leaves) into pond mesocosms would increase reciprocal export (frog biomass) to the surrounding terrestrial ecosystem. In contrast, we found that increases in primary productivity consistently decreased frog biomass, except with leaf litter inputs. With leaf inputs, primary productivity did not affect the export of frogs, indicating that leaf detritus and associated microbial communities may be more important than algae for frog production. We found that subsidy inputs tended to increase reciprocal exports, and thus partial concordance with our conceptual framework.