Spatiotemporal allozyme variation in the damselfly, Lestes viridis (Odonata: Zygoptera): gene flow among permanent and temporary ponds

Several insect species seem to persist not only in permanent but also in temporary ponds where they face particularly harsh conditions and frequent extinctions. Under such conditions, gene flow may prevent local adaptation to temporary ponds and may promote phenotypic plasticity, or maintain apparent population persistence. The few empirical studies on insects suggest the latter mechanism, but no studies so far quantified gene flow including both pond types. We investigated the effects of pond type and temporal variation on population genetic differentiation and gene flow in the damselfly Lestes viridis in northern Belgium. We report a survey of two allozyme loci (Gpi, Pgm) with polyacrylamide gel electrophoresis in 14 populations from permanent and temporary ponds, and compared these results with similar data from the same permanent populations one year before. The data suggested that neither pond-drying regime, nor temporal variation have a substantial effect on population genetic structuring and did not provide evidence for stable population differentiation in L. viridis in northern Belgium. Gene flow estimates were high within permanent and temporary ponds, and between pond types. Our data are consistent with a source-sink metapopulation system where temporary ponds act as sinks in dry years, and are quickly recolonized after local population extinction. This may create a pattern of apparent population persistence of this species in permanent and temporary ponds without clear local adaptation.     

Pond tadpoles with generalized morphology: is it time to reconsider their functional roles in aquatic communities?

With rare exceptions, anuran larvae have traditionally been considered to occupy lower trophic levels in aquatic communities where they function as microphagous suspension feeders. This view is being challenged by studies showing that tadpoles with generalized morphology often function as macrophagous predators. Here, we review the literature concerning macrophagy by tadpoles and provide two additional examples involving generalized tadpoles. In the first, we demonstrate with laboratory and field experiments that wood frog (Rana sylvatica) tadpoles are major predators of macroinvertebrates in ponds. In the second, we show that green frog (R. clamitans) tadpoles can cause catastrophic reproductive failure of the wood frog via egg predation. These results and data from other studies challenge the assumption that generalized tadpoles function as filter-feeding omnivores, and question the general applicability of community organization models which assume that predation risk increases with pond permanence. We suggest that predation risk is greater in temporary ponds than in more permanent ponds for many organisms that are vulnerable to predation by tadpoles. This being so, a conditional model based upon interactions that are species-specific, life-stage-specific, and context-dependent may better explain community organization along hydrological gradients than models which assume that temporary ponds have few or no predators. Received: 30 November 1998 / Accepted: 2 May 1999     

Crustacean zooplankton community structure in temporary and permanent grassland ponds

Lentic community structure varies across a size gradient of ponds and lakes with physical factors, such as pond drying, and biotic factors, such as fish predation, determining the species assemblage. We studied the effects of pond drying and fish absence on crustacean zooplankton across a gradient of pond sizes in a Texas grassland. We determined the species compositions and size distributions of crustacean zooplankton in 20 temporary and 18 permanent ponds in April after March rains had refilled the ponds. The surface areas of temporary and permanent ponds ranged from <0.01 to 0.21 ha and 0.04 to 13.8 ha, respectively, and temporary ponds were significantly smaller, on average, than permanent ponds. Fish were absent from all temporary ponds and present in all permanent ponds. We detected a difference in the zooplankton species assemblages of the temporary and permanent ponds. Out of 14 total zooplankton taxa that occurred in eight or more ponds, seven taxa were significantly more prevalent in temporary ponds and four taxa were significantly more prevalent in permanent ponds. The sizes of zooplankton in the temporary fishless ponds were greater than those in the permanent ponds with fish present. We concluded that pond size mediated susceptibility to pond drying, and pond drying determined the presence and absence of fish and their secondary trophic-level effect on zooplankton community structure. Handling editor: Steven Declerck     

Morphological and life history responses to predators in larvae of seven anurans

Amphibian larvae often face two major sources of mortality: pond desiccation and predation. Tadpoles of seven anuran species with different preferences for type of breeding habitat, on a hydroperiod scale, were tested for responses to the presence of predators by raising them experimentally in the presence and absence of a separately caged invertebrate predator that was fed on conspecific tadpoles. The species typically breeding in temporary or semi-permanent ponds ( Rana temporaria , Rana arvalis , Rana dalmatina and Hyla arborea ) – where invertebrate predator populations are predicted to vary considerably spatiotemporally – all showed marked induced increases in tail fin depth in response to predator presence. These species also tended to respond by reduced growth rates. The representative of the most ephemeral habitats, Bufo calamita , did not respond in any of these traits. Its congeneric, Bufo bufo , a toxic inhabitant of permanent ponds and lakes, tended to respond to predator presence by reducing its growth rate, though not by a tail depth increase. I argue that the rather poor swimming performance in Bufo tadpoles may opt for defences other than locomotor ability. The palatable, permanent pond species Pelobates fuscus did not alter either its growth rate or tail morphology. Possible explanations for this result are discussed.     

Diurnal habitat segregation by tadpoles in two temporary ponds in an Atlantic Rainforest remnant, Northeastern Brazil

Tadpoles are found in diverse aquatic habitats, where they share resources in various dimensions. Tadpole assemblages were studied in two temporary ponds in Atlantic Rainforest. Our aim was to analyze the temporal and spatial distribution patterns and their correlation with abiotic factors. Tadpoles were sampled weekly using nets, from March to December 2008. Samples were collected during the day and at night, and taken near the shore and in the centre of the ponds. Samples in inshore and pond centres were taken from both areas with and without aquatic vegetation. The assemblage consisted of 15 species in pond A and 12 species in pond B. The majority of species was accidental, and used the pond continuously. The assemblage was predominantly diurnal, which may be related to greater activity rate. In pond A the tadpoles primarily used the centre of ponds, whereas in pond B tadpoles primarily used shore areas. However, some populations showed preferences for particular environments. In pond A, environmental heterogeneity in the centre had the greatest influence on tadpole abundance. Our data suggest that temporal distribution facilitated the coexistence of different species of tadpoles in the pond but this conclusion is tentative and requires verification. Hence, more data needs to be collected so that we can propose effective area management plans.     

Evaluating predation pressure on green treefrog larvae across a habitat gradient

The effect of a predator on the abundance of a prey species depends upon the predators abundance and its ability to capture that prey. The objectives of this research were to evaluate the community structure of predators of green treefrog (Hyla cinerea) tadpoles across habitat types and evaluate the effectiveness of individual predators on H. cinerea tadpoles. Correspondence and cluster analyses of predator frequencies across 23 aquatic habitats indicated that the majority of variance in predator communities was due to a division between permanent and temporary habitats. Experimental work demonstrated that survival of the smallest H. cinerea tadpoles was significantly lower than survival of medium and large tadpoles with the most effective predators, indicating that H. cinerea tadpoles attain a refuge from predation at larger body sizes. We combined the effectiveness of predators in experiments with the abundance of each predator species from the predator community survey to demonstrate that predation pressure on H. cinerea tadpoles is higher in temporary ponds. This pattern may explain in part why this species generally breeds successfully only in permanent habitats. It also confirms that discussions about an increasing gradient of predation pressure from temporary to permanent aquatic habitats should be restricted to individual prey species for which such a gradient has been demonstrated.     

Spatiotemporal allozyme variation in the damselfly, <Emphasis Type="BoldItalic">Lestes viridis</Emphasis> (Odonata: Zygoptera): gene flow among permanent and temporary ponds

Several insect species seem to persist not only in permanent but also in temporary ponds where they face particularly harsh conditions and frequent extinctions. Under such conditions, gene flow may prevent local adaptation to temporary ponds and may promote phenotypic plasticity, or maintain apparent population persistence. The few empirical studies on insects suggest the latter mechanism, but no studies so far quantified gene flow including both pond types. We investigated the effects of pond type and temporal variation on population genetic differentiation and gene flow in the damselfly Lestes viridis in northern Belgium. We report a survey of two allozyme loci (Gpi, Pgm) with polyacrylamide gel electrophoresis in 14 populations from permanent and temporary ponds, and compared these results with similar data from the same permanent populations one year before. The data suggested that neither pond-drying regime, nor temporal variation have a substantial effect on population genetic structuring and did not provide evidence for stable population differentiation in L. viridis in northern Belgium. Gene flow estimates were high within permanent and temporary ponds, and between pond types. Our data are consistent with a source-sink metapopulation system where temporary ponds act as sinks in dry years, and are quickly recolonized after local population extinction. This may create a pattern of apparent population persistence of this species in permanent and temporary ponds without clear local adaptation.     

Competing tadpoles: Australian native frogs affect invasive cane toads (Rhinella marina) in natural waterbodies

The cane toad (Rhinella marina) is one of the most successful invasive species worldwide, and has caused significant negative impacts on Australian fauna. Experimental work in the laboratory and in mesocosms has shown that tadpoles of native frogs can affect survival, size at metamorphosis and duration of larval period of cane toad tadpoles. To test if these effects occur in nature, we conducted a field experiment using two temporary ponds where we set up enclosures with tadpoles of native green tree frogs (Litoria caerulea) and cane toads in treatments with a range of densities and combinations. The presence of green tree frog tadpoles significantly decreased the growth rate of toad tadpoles and increased the duration of their larval period in both ponds; in one pond, frog tadpoles also significantly reduced the body length and mass of metamorph toads. Toad tadpoles did not have any significant negative effects on green tree frog tadpoles, but there was strong intraspecific competition within the latter species: increased frog tadpole density resulted in increased larval period and reduced survival, growth rate and size at metamorphosis for frogs at one or both ponds. Our results are encouraging for the possibility of using native frogs as one component of an integrated approach to the biological control of cane toads.     

Canopy Closure,Competition, and the Endangered Dusky Gopher Frog

Abstract: A major challenge facing wildlife biologists is understanding why some species go extinct while others persist in the same habitat. To address this question, we investigated whether tree canopy closure over ponds affects growth and survival of rare and common tadpoles within ponds and mediates competitive interactions among species. We conducted 2 experiments to test whether canopy closure and competition may have contributed to the decline of the endangered dusky gopher frog (Rana sevosa), but allowed the persistence of the southern leopard frog (R. sphenocephala). We explored the response of both species to canopy closure in single-species and mixed- (1:1) species treatments of identical total tadpole density. An experiment using aquatic enclosures in temporary ponds showed that canopy closure reduced tadpole growth approximately 20% for both species. Survival of dusky gopher frog tadpoles was higher in mixed-species enclosures than in single-species enclosures. In a complementary experiment using artificial ponds, dusky gopher frogs had lower survival to metamorphosis, reduced size at metamorphosis, and produced a lower total biomass of metamorphosed juveniles in shaded ponds. Southern leopard frogs exhibited reduced body size at metamorphosis only when shaded. These studies suggest that pond canopy closure, not larval competition, may be contributing to the decline of the dusky gopher frog. The different responses to canopy closure suggest a potential mechanism for the loss of dusky gopher frogs and the persistence of southern leopard frogs. Removal of trees from historically open-canopy ponds may help facilitate the recovery of dusky gopher frogs and benefit similar species.     

Invertebrate communities and turnover in wetland ponds affected by drought

1. Changes to the macroinvertebrate fauna found in small ponds on a freshwater marsh (Aberlady Bay, Scotland) in 1986, 1987 and 1992 were used to assess the impact of the 1988–92 drought on taxon turnover in pond communities. 2. Permanent ponds accumulated taxa over the study period. Ponds that were wet throughout 1986–87 but dried in 1992 lost some of the many taxa associated with permanent water but acquired a reduced fauna typical of temporary ponds. Ponds that were temporary in 1986–87 were dry during most of 1992 and lost almost all aquatic taxa. 3. Faunal turnover was considerable even in permanent ponds. Extinction rates were high for taxa typical of permanent or temporary ponds. Colonization rates were poor for the taxa from permanent water, but high for the taxa from temporary ponds. 4. The importance of colonization and extinction rates as main predictors of the distribution of species between the ponds was looked at using metapopulation incidence functions, where observed incidence of a taxon can be used to predict colonization and extinction rates. Predicted rates were then compared with observed rates. Incidence functions gave reasonable predictions of observed colonization rates but were poor predictors of extinction, even for taxa that appeared likely to be true metapopulations. 5. For the pond fauna, including fugitive species adapted to temporary ponds, whilst colonization may well depend on environmental stochasricity (how long a pond holds water), subsequent survival depends on other, demographic, processes (e.g. finding a mate, predation) rather than the pond drying out.     

Predation and the distribution and abundance of a pulmonate pond snail

Summary The abundances of a freshwater pulmonate snail, Lymnaea elodes were studied in a temporary pond and a permanent, more productive pond in northeastern Indiana, USA. When snails from both populations were reared in each of the ponds in containers excluding predators, snails grew to be 1.3 to 2 times as large in the more productive pond, and laid 9 times as many eggs. However, field sampling data showed adults to be more abundant in the temporary pond. The only obvious difference between the two ponds was the presence of the molluscivorous central mudminnow (Umbra limi) in the permanent pond. These fish fed upon L. elodes when eggs and juvenile snails were abundant. In an experiment in the temporary pond, addition of mudminnows lowered egg and juvenile snail survival in pens where snail abundances had been increased. We suggest that vertebrate predators like the mudminnow can be significant sources of mortality for thin shelled species like L. elodes, possibly excluding them from habitats like lakes and rivers.Address for offprint requests     

Foraging tactics in alternative heterochronic salamander morphs: trophic quality of ponds matters more than water permanency

1. In lentic freshwater habitats, the composition of animal assemblages shifts along a gradient from temporary to permanent basins. When habitats with different degrees of permanence are at the scale of the home range of species, they constitute alternatives in terms of energy acquisition through feeding. 2. In this context, previous studies showed an advantage of metamorphic over paedomorphic tiger salamanders (Ambystoma tigrinum) in temporary ponds which are only available to metamorphs. The aim of this study was to establish whether salamanders obtain similar benefits in ponds that do not differ in water permanence and whether salamanders shifted from detrimental to advantageous ponds. To this end, we determined the feeding habits, body condition and movement patterns of the two morphs in a complex of four permanent and four temporary ponds. 3. Consistent with previous studies, metamorphs consumed higher‐quality diets than paedomorphs in term of energy intake. However, these differences occurred because metamorphs consumed fairy shrimp in a single temporary pond. Individual movement patterns confirmed that most of the metamorphs used different aquatic habitats both within and between years and that most of them moved from permanent ponds for breeding towards the most profitable temporary pond in terms of foraging. 4. These results indicate that habitat selection by salamanders is optimal in term of energy intake in metamorphs that use high quality ponds independently of hydroperiod. It seems that both spatial and temporal variation can influence the relative foraging success of each morph.     

Relating fish kills to upwellings and wind patterns in the Salton Sea

Awareness of pond conservation value is growing all over Europe. Ponds are recognized as important ecosystems supporting large numbers of species and several rare and threatened aquatic plants, macroinvertebrates and amphibians. Notwithstanding ponds, particularly temporary ones, are still neglected in Italy. There are some gaps in our understanding of the macrophyte ecology and the conservation value of Mediterranean small still waters. Therefore, this study investigated the macrophyte communities and physico-chemical characteristics of 8 permanent and 13 temporary ponds along the Tyrrhenian coast near Rome, with the aim to relate the distribution of aquatic plants to environmental variables, and to define the botanical conservation value of ponds. Throughout the study period (Spring 2002), Principal Component Analysis performed on abiotic variables clearly discriminated temporary ponds, smaller and more eutrophic, from permanent ponds, larger and with higher pH and oxygen concentration. A total of 73 macrophyte taxa were collected in the study ponds. Temporary waters hosted a smaller number of plant species than permanent ones. Besides hydroperiod length, the environmental factors related to plant richness were maximum depth, surface area, dissolved oxygen and nitrogen concentration in the water. Moreover, the Non-metric Multidimensional Scaling showed a high dissimilarity in the taxonomic composition of aquatic plants between temporary and permanent ponds. The former contained more annual fast-growing species (Callitriche sp. pl. and Ranunculus sp. pl.), while in the latter species with long life-cycles (i.e. Potamogeton sp. pl.) were more abundant. Our results highlighted that temporary and permanent ponds in central Italy have different macrophyte assemblages, with aquatic species (including some of conservation interest at regional scale) exclusively found in each pond type. This suggested that both type of ponds could give an irreplaceable contribution to the conservation of aquatic plant diversity of these freshwater ecosystems. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

Predator‐induced phenotypic plasticity in an arid‐adapted tropical tadpole

Adaptive phenotypic plasticity is widespread and involves diverse phenotypes. Key environmental stressors, such as predation risk, can simultaneously induce changes in multiple traits, but the magnitude of response is dependent upon the environmental conditions. Species that utilize temporary ponds are expected to exhibit stronger predator‐induced responses in the form of morphology than behaviour (i.e. reduced activity) to meet the demands of rapid development by maintaining high foraging activity while reducing predation risk via morphologically plastic traits. In a laboratory experiment, I examined the effects of predator chemical cues and conspecific alarm cues on activity, development and morphology on Leptodactylus bufonius tadpoles. This species has terrestrial oviposition and completes the early part of its development outside of ephemeral and temporary ponds in the Gran Chaco ecoregion of South America. Tadpoles in the predator treatments exhibited both behavioural and morphological predator‐induced plastic responses. Tadpoles tended to possess shorter, deeper tails when exposed to predators. The greatest reduction in activity was observed in tadpoles exposed to both predator and conspecific alarm cues, which subsequently resulted in the slowest development. Temporary and ephemeral pond adapted species with terrestrial oviposition may capitalize on a head start in development by being able to afford reduced growth rates via a reduction in activity. This may occur when the constraints imposed by pond hydroperiod (e.g. risk of pond drying) are relaxed when compared with species with aquatic oviposition, which must undergo all stages of development during the pond's hydroperiod. Thus, in addition to the predator and hydroperiod gradients, examining phenotypically plastic responses along a ‘terrestriality gradient’ in a comparative framework would provide insights as to the costs and benefits of increasing terrestriality in anuran reproductive modes to environmental stressors.     

Trait flexibility of generalist invertebrates exposed to contrasting predation and drying stressors

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Evaluating a predator–prey interaction in the field: the interaction between beetle larvae (predator) and tadpoles (prey)

The larval amphibian community of temporary pond ecosystems has served as a model for studies in community ecology, with a majority of this work being conducted in mesocosms. Recent research has suggested that mesocosms may overestimate ecological effects; therefore, experimental studies conducted under field conditions are required to gauge the results of mesocosm studies. To assess a species interaction under more natural conditions, we conducted a series of field experiments examining the predator–prey interaction between beetle larvae ( Dytiscus sp.; predator) and larval wood frogs Rana sylvatica (prey) in central Pennsylvania, USA. Quantitative sampling of woodland ponds indicated that beetle larvae of the genus Dytiscus were the most common predator of tadpoles. In a field enclosure experiment, dytiscids were effective predators of tadpoles in the pond environment. Moreover, tadpoles avoided areas in a pond containing caged dytiscids, demonstrating that tadpoles recognize the chemical stimuli of predators in complex environments. The results of this study are consistent with data from prior laboratory and mesocosm studies and suggest that these venues can produce reliable interpretations of predator–prey dynamics in this community.     

PERFORMANCE OF TADPOLES FROM THE HYBRIDOGENETIC RANA ESCULENTA COMPLEX: INTERACTIONS WITH POND DRYING AND INTERSPECIFIC COMPETITION

The performance of three genotypes (LL, LR, RR) of tadpoles resulting from the hybrid mating system of Rana lessonae (phenotype L, genotype LL) and Rana esculenta (phenotype E, genotype LR) was determined in artificial ponds. The effects of interspecific competition and pond drying on growth, development, and survival of tadpoles were used to measure the performance of genotypes and the relative fitness of offspring. Among the three genotypes, tadpoles from the homogametic mating RR had the lowest survival, growth, and development under all environmental conditions. Body size of the LL and LR genotype tadpoles at metamorphosis was reduced by competition and pond drying. Days to metamorphosis were also higher for the LL and LR genotype tadpoles in competition ponds. The proportion of individuals metamorphosing of each genotype was differentially lowered by competition and pond drying. The LL genotype produced more metamorphs than the LR genotype in the constant water level ponds, but the LR genotype produced more in drying ponds. In competition ponds, the LR genotype produced more metamorphs than the LL genotype, but the LL genotype produced more metamorphs in ponds without competition. The RR genotype produced no metamorphs in any of the experimental environments. Increased performance of LR offspring from the heterogametic mating, in harsh conditions, and reduced performance of RR offspring from the homogametic mating, even under favorable conditions, relative to the parental genotype (LL) suggests that the population dynamics of this hybridogenetic system is strongly dependent on mate choice in mixed populations and the subsequent pond environment females select for oviposition and larval development.     

The role of historic and climatic factors in the distribution of crustacean communities in Iberian Mediterranean ponds

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Synergistic effects of multiple mechanisms drive priority effects within a tadpole assemblage

Mounting evidence suggests that the history of species arrival to a locality can have important effects on species performance but the mechanism(s) through which priority effects are produced is not always clear. Differences in the developmental time of frog tadpoles provide an opportunity to examine mechanisms through which priority effects may influence fitness components of a late arriving taxon. Specifically, tadpoles of the southern leopard frog (Rana) can often require more than a year to complete metamorphosis so they overwinter in a pond and compete with newly deposited tadpoles in the spring. We conducted an experiment in artificial ponds to evaluate mechanisms through which overwintering Rana tadpoles influence fitness components of southern toad (Bufo) tadpoles deposited into ponds during the spring. We found that overwintered Rana reduced Bufo performance while newly hatched Rana that enter a pond simultaneously with Bufo did not. The production of this priority effect was primarily the result of Rana depleting algal resources in a pond during the winter prior to Bufo arrival. The performance of Bufo did not correspond with variation in the abundance of algae present in a pond during the spring and we found evidence to indicate that, in the absence of resource exploitation during the winter, overwintered Rana do not compete strongly with Bufo during the spring when both species co‐occur. When Rana deplete algal resources during the winter, however, interactions between Rana and Bufo during the spring became much more important as both the development rate and survival of Bufo was reduced to a greater extent than what would have been predicted by resource exploitation alone. Our results demonstrate that priority effects can result from early colonists depleting resource availability prior to the arrival of other species which can intensify behavioral/physical interactions between species when they co‐occur.