Growth and foraging consequences of facultative paedomorphosis in the tiger salamander,Ambystoma tigrinum nebulosum

Summary Facultative paedomorphosis in salamanders occurs when larvae respond to varying environmental conditions by either metamorphosing into terrestrial metamorphic adults or retaining their larval morphology to become sexually mature paedomorphic adults. Several hypotheses have been proposed for the evolutionary maintenance of this environmentally induced dimorphism, but few data are available to assess them adequately. We studied a montane population of the tiger salamander,Ambystoma tigrinum nebulosum, and measured the adult growth rate and body condition across three growing seasons to assess the relative costs and benefits of each morph. Metamorphic adults grew more than paedomorphic adults in terms of snout—vent length across years and in weight within years. Dietary analyses and foraging experiments revealed some of the proximate factors that may underlie these differential growth patterns. Across all prey, metamorphs had significantly higher biomass and calories per stomach sample than paedomorphs. Metamorphic diets primarily consisted of the fairy shrimpBranchinecta coloradensis, whereas paedomorphic diets contained a variety of benthic and terrestrial invertebrates. Foraging experiments revealed that both morphs are more successful at capturing fairy shrimp relative to other prey types and both show high electivity toward this prey. However, fairy shrimp occurred only in non-permanent ponds and thus are inaccessible to paedomorphs, which can survive only in permanent ponds. Paedomorphs also experience higher levels of intraspecific competition with large larvae in permanent ponds than metamorphs do in non-permanent ponds. Thus, metamorphs obtain a growth advantage over paedomorphs by foraging in non-permanent ponds that contain fairy shrimp and have reduced intraspecific competition. These results suggest that paedomorphs should have decreased fitness relative to metamorphs, primarily because metamorphs can move into the best habitats for growth. The net fitness effect of morph-specific differences in dispersal depend on whether there are trade-offs with other life history traits. Nonetheless, because the relative benefit of metamorph dispersal will change with environmental conditions in permanent ponds and the surrounding habitat, the relative fitness payoff to each morph should track changes in the environment. Thus, facultative paedomorphosis may be maintained in part by variable, environmentally-specific fitness payoffs to each morph.     

Caddisfly life histories along permanence gradients in high-altitude wetlands in Colorado (U.S.A.)

SUMMARY 1. Larvae of cased caddisflies (Limnephilidae and Phryganeidae) are among the most abundant and conspicuous invertebrates in northern wetlands. Although species replacements are often observed along permanence gradients, the underlying causal mechanisms are poorly understood. In this paper, we report on the distributional patterns of caddisflies in permanent and temporary high‐altitude ponds, and how those patterns reflect differences in life history characteristics that affect desiccation tolerance (fundamental niches) versus constraints related to biotic interactions (realised niches). 2. Species (Hesperophylax occidentalis and Agrypnia deflata) that were encountered only in permanent ponds are restricted in distribution by life history (no ovarian diapause, aquatic oviposition, and/or inability to tolerate desiccation). Although the egg masses of H. occidentalis tolerate desiccation, the larvae leave the protective gelatinous matrix of the egg mass because adults oviposit in water. 3. Three species (Asynarchus nigriculus, Limnephilus externus and L. picturatus) have life history characteristics (rapid larval growth, ovarian diapause and terrestrial oviposition of desiccation‐tolerant eggs) that should facilitate the use of both permanent and temporary habitats. However, A. nigriculus is rare or absent in most permanent ponds, and L. externus and L. picturatus are rare or absent in most temporary ponds. Experimental data from a previous study on the combined effects of salamander predation and interspecific interactions among caddisflies (e.g. intraguild predation) suggest that biotic interactions limit each species to a subset of potentially exploitable habitats. 4. Many wetland invertebrates exhibit species replacements along permanence gradients, but few studies have separated the relative importance of the effects of drying per se from the effects of biotic interactions. Our results emphasise the complementary roles of comparative data on life histories and experimental data on competition and predation for understanding invertebrate distributions along permanence gradients.     

Linking the evolution of habitat choice to ecosystem functioning: direct and indirect effects of pond-reproducing fire salamanders on aquatic-terrestrial subsidies

Shifts in life history traits and in the behaviour of species can potentially alter ecosystem functioning. The reproduction of the central European fire salamander (Salamandra salamandra), which usually deposits its larvae in first-order streams, in small pool and pond-like habitats, is an example of a recent local adaptation in this species. Here we aimed to quantify the direct and indirect effects of the predatory larvae on the aquatic food webs in the ponds and on the flux of matter between the ponds and adjacent terrestrial habitats. Our estimates are based on biomass data of the present pond fauna as well as on the analysis of stomach content data, growth rates and population dynamics of the salamander larvae in pond habitats. By their deposition of larvae in early spring, female fire salamanders import between 0.07 and 2.86 g dry mass m^?2 larval biomass into the ponds. Due to high mortality rates in the larval phase and the relatively small size at metamorphosis of the pond-adapted salamanders compared to stream-adapted ones, the biomass export of the metamorphosed salamanders clearly falls below the initial biomass import. Catastrophic events such as high water temperatures and low oxygen levels may even occasionally result in mass mortalities of salamander larvae and thus in a net 100 % import of the salamander biomass into the pond food webs. Indirect effects further accelerate this net import of matter into the aquatic habitat, e.g. the feeding of salamanders on aquatic insect larvae with the emergence of terrestrial adults—thus preventing export—and on terrestrial organisms that fall on the water surface (supporting import). This study demonstrates that the adaptation of salamanders to pond reproduction can alter food web linkages across ecosystem boundaries by enhancing the flux of materials and energy from terrestrial (i.e. forest) to the aquatic (i.e. pond) habitat.     

Growth and population structure of Salamandra salamandra (L.) larvae in different limnological conditions

Salamandra salamandra (L.) larvae were studied in three types of aquatic systems, a permanent spring, a rock pool and temporary ponds. Growth was followed throughout ontogenesis until completion of metamorphosis. Growth patterns changed according to the pond type. Temporary ponds were rich in organic matter and subsequently larvae matured more rapidly there than those in a permanent pool or a spring. In the latter, cannibalism appears to be of major ecological importance, enabling survival and metamorphosis of large size tadpoles. The pattern of growth and metamorphosis is compared with European salamanders and the differences are discussed considering adaptations for survival of this species under xeric conditions.     

Effects of hydroperiod and predation on a Mississippi River floodplain invertebrate community

 The objective of this study was to determine if pond permanence and vertebrate predation (by fish and waterfowl) affect invertebrate community structure in the mudflat habitat of floodplain ponds. Invertebrate communities were studied for 1 year in four Mississippi River floodplain ponds with different hydroperiods. Pond 1 experienced five dry periods, pond 2 experienced four, pond 3 dried once, and standing water remained in pond 4 for the entire year. Vertebrate predator exclusion treatments (all access, no access, small-fish access and cage controls) were placed in all ponds. As pond duration increased, predatory invertebrate richness and abundance increased while overall invertebrate richness and abundance decreased. With the exception of the cladoceran Diaphanosoma, all commonly encountered taxa were strongly affected by pond permanence in terms of abundance, biomass and, generally, individual biomass. Taxa were nearly early divided between those that were more abundant in less permanent ponds and those that were more abundant in longer-duration ponds. Invertebrate taxa richness, abundance, and total biomass were lower in the all-access treatment than in the treatments that restricted predator access, and these effects were stronger in the more permanent ponds. In general, there were no significant differences in responses to the treatments with small-fish access and no access. These results support models that predict relatively weak effects of predation in frequently disturbed habitats. Received: 30 May 1995 / Accepted: 21 June 1996     

Prioritizing Conservation of Coastal Ponds for Wintering Redheads

The Laguna Madres of Texas, USA, and Tamaulipas, Mexico, are the most important wintering areas for redheads (Aythya americana) as most of the continental population winters in these lagoons. Redheads forage in the saline waters of the Laguna Madre and make daily flights to coastal freshwater ponds on the adjacent mainland to drink. The abundance and spatial distribution of coastal ponds varies depending on precipitation and can influence the foraging pressure on adjacent seagrass meadows. We conducted weekly aerial surveys to monitor coastal pond use by wintering redheads from mid-October through mid-March along the entire length of the Laguna Madre of Texas, during 2000–2003 and in 2012–2014. We developed 3 parameters to provide a measure of biological value of each coastal pond to redheads: amount of foraging habitat within 10 km of each pond, water permanence of the pond, and the potential to distribute redheads if inundated. During 101 aerial surveys across 5 years of study, we identified 140 coastal ponds that were used by redheads. We developed a prioritization scheme to identify wetlands that remain inundated in all years and targeted them for conservation protection. We identified those coastal ponds that, if enhanced through increasing their water permanence, would provide additional drinking sites during dry years and help distribute redheads on more foraging habitat, thereby reducing potential overgrazing on seagrass meadows. We identified 3,624 ha of foraging habitat (21.5% of all foraging habitat) in the lower Laguna Madre that had no coastal ponds within a 10-km radius and, thus, was proximal to potential areas for coastal pond creation. Our results provide guidance for resource managers to protect, enhance, or create coastal ponds to reduce foraging pressure on seagrass meadows in the Laguna Madre and help sustain future populations of wintering redheads. © 2021 The Wildlife Society.     

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     

Does interspecific competition drive patterns of habitat use in desert bat communities?

Bodies of water are a key foraging habitat for insectivorous bats. Since water is a scarce and limiting resource in arid environments, bodies of open water may have a structuring effect on desert bat communities, resulting in temporal or spatial partitioning of bat activity. Using acoustic monitoring, we studied the spatial and temporal activity patterns of insectivorous bats over desert ponds, and hypothesised that sympatric bat species partition the foraging space above ponds based on interspecific competitive interactions. We used indirect measures of competition (niche overlap and competition coefficients from the regression method) and tested for differences in pond habitat selection and peak activity time over ponds. We examined the effect of changes in the activity of bat species on their potential competitors. We found that interspecific competition affects bat community structure and activity patterns. Competing species partitioned their use of ponds spatially, whereby each species was associated with different pond size and hydroperiod (the number of months a pond holds water) categories, as well as temporally, whereby their activity peaked at different hours of the night. The drying out of temporary ponds increased temporal partitioning over permanent ponds. Differences in the activity of species over ponds in response to the presence or absence of their competitors lend further support to the role of interspecific competition in structuring desert bat communities. We suggest that habitat use and night activity pattern of insectivorous bats in arid environments reflect the trade-offs between selection of preferred pond type or activity time and constraints posed by competitive interactions.     

Predator defense along a permanence gradient: roles of case structure, behavior, and developmental phenology in caddisflies

Species replacements along freshwater permanence gradients are well documented, but underlying mechanisms are poorly understood for most taxa. In subalpine wetlands in Colorado, the relative abundance of caddisfly larvae shifts from temporary to permanent basins. Predators on caddisflies also shift along this gradient; salamanders (Ambystoma tigrinum nebulosum) in permanent ponds are replaced by predaceous diving beetles (Dytiscus dauricus) in temporary habitats. We conducted laboratory and field experiments to determine the effectiveness of caddisfly cases in reducing vulnerability to these predators. We found that larvae of a temporary-habitat caddisfly (Asynarchus nigriculus) were the most vulnerable to salamanders. Two relatively invulnerable species (Limnephilus externus, L. picturatus) exhibited behaviors that reduced the likelihood of detection and attack, whereas the least vulnerable species (Agrypnia deflata) was frequently detected and attacked, but rarely captured because cases provided an effective refuge. Vulnerability to beetle predation was also affected by cases. The stout cases of L. externus larvae frequently deterred beetle larvae, whereas the tubular cases of the other species were relatively ineffective. Two of these vulnerable species (A. nigriculus and L. picturatus) often co-occur with beetles; thus, case construction alone is insufficient to explain patterns of caddisfly coexistence along the permanence gradient. One explanation for the coexistence of these two species with beetles is that they develop rapidly during early summer and pupate before beetle larvae become abundant. One species (L. picturatus) pupates by burying into soft substrates that serve as a refuge. The other (A. nigriculus) builds stone pupal cases, which in field experiments, more than doubles survival compared to organic pupal cases. The combined results of these experiments suggest that caddisfly distributions along permanence gradients depend on a suite of primary and secondary predator defenses that include larval and pupal case structure, predator-specific escape behaviors, and the phenology of larval development.     

Tadpole competition in a desert anuran community

Summary Five interspecific competition experiments performed in artificial ponds under temporary pond-like conditions established that temporary and permanent pond tadpoles interact. Temporary pond species typically lowered survivorship and reduced progress towards metamorphosis of permanent pond tadpoles. These results suggest that competitive interactions may partially determine the local distribution of tadpoles in the Chihuahuan Desert by preventing some permanent pond anurans from using temporary ponds. Predation by Scaphiopus multiplicatus tadpoles (a temporary pond species) may also prevent permanent pond tadpoles from using temporary ponds.     

Rana temporaria metamorph production and population dynamics in the field – Effects of tadpole density,predation and pond drying

This study investigates factors of importance for tadpoles survival and metamorph production in the common frog Rana temporaria. It also assess the importance of this for the population dynamics of the species. Eighteen ponds were studied for up to 8 years. Data collected each year included: number of spawn clumps deposited, tadpole number and metamorph number. The permanency of the ponds was also recorded each year. Measures were taken of predator density. There was no suggestion of density dependence in the survival of tadpoles. In contrast, the number of spawn clumps deposited per pond area was highest for ponds with high survival. Density of predators (sticklebacks, newts and invertebrates) was negatively correlated to tadpole and metamorph survival. This was true both within (among years) and among ponds. Several of the study ponds dried completely before metamorphosis in some years. However, those ponds also were those with the smallest number of predators and in years with successful metamorphosis, these ponds produced more metamorphs than more permanent ponds. An analysis of the year to year dynamics showed that population size (number of deposited spawn clumps) was correlated to that in the previous year, suggesting a fairly high adult survival, but also on the number of metamorphs emerging two or three years before (corresponding to the age of sexual maturity of the species). It is concluded that the aquatic stage is not strongly limiting in these ponds but conservation efforts should be focused on the terrestrial habitat. Also, the study stresses the value of temporary ponds, despite the fact that recruitment often fails totally in these.     

Are cladoceran diversity and community structure linked to spatial heterogeneity in urban landscapes and pond environments?

Biodiversity patterns in cladoceran communities were investigated in urban waterbodies in relation with residential land use, pond management, and waterbody environments. We evaluated species richness in the pelagic and littoral zones of eighteen waterbodies of a large Canadian city. Gamma diversity (26 species) observed at a small scale in the urban survey was important comparatively to large-scale surveys of lakes. Beta diversity ranged from 1 to 8 species among waterbodies. We tested if littoral species greatly contributed to regional diversity in urban waterbodies. Littoral species (Chydoridae, Ilyocryptidae, Macrothricidae, Polyphemidae) accounted for 58% of the total species pool. We distinguished five cladoceran assemblages associated to different waterbodies (temporary ponds, permanent lakes, and wetlands). Cladoceran communities were more diverse and variable in permanent lakes than in temporary ponds. Changes in cladoceran species assemblages among waterbodies were driven by variations in waterbody size and phosphorus enrichment, macrophyte and algal biomass, urban density, pond management practices, and the presence of potential predators as fish and macroinvertebrates. Our study indicates that both artificial ponds and lakes and natural wetlands are valuable habitats for the conservation of cladoceran biodiversity and rare endemic species in urban regions. Further research on pond management strategies promoting urban aquatic biodiversity should be undertaken.     

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

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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     

Predator size and phenology shape prey survival in temporary ponds

Theoretical efforts suggest that the relative sizes of predators and their prey can shape community dynamics, the structure of food webs, and the evolution of life histories. However, much of this work has assumed static predator and prey body sizes. The timing of recruitment and the growth patterns of both predator and prey have the potential to modify the strength of predator–prey interactions. In this study, I examined how predator size dynamics in 40 temporary ponds over a 3-year period affected the survival of spotted salamander (Ambystoma maculatum) larvae. Across communities, gape-limited predator richness, but not size, was correlated with habitat duration (pond permanence). Within communities, mean gape-limited predator size diminished as the growing season progressed. This size reduction occurred because prey individuals grew into a body size refuge and because the largest of the predators left ponds by mid-season. Elevated gape-limited predation risk across time and space was predicted by the occurrence of two large predatory salamanders: marbled salamander larvae (Ambystoma opacum) and red-spotted newt adults (Notophthalmus viridescens). The presence of the largest gape-limited predator, A. opacum, predicted A. maculatum larval survival in the field. The distribution of large predatory salamanders among ponds and across time is expected to lead to differing community dynamics and to generate divergent natural selection on early growth and body size in A. maculatum. In general, a dynamic perspective on predator size often will be necessary to understand the ecology and evolution of species interactions. This will be especially true in frequently disturbed or seasonal habitats where phenology and ontogeny interact to determine body size asymmetries. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Landscape connectivity affects individual survival in unstable patch networks: The case of a freshwater turtle inhabiting temporary ponds

1. The loss of connectivity is among the main threats for species occupying freshwater pond networks. Landscape connectivity can impact the persistence of patchy populations by reducing movement rates among ponds, thereby increasing the likelihood of local extinctions in source–sink systems, and reducing the probability of colonisation following extinctions. In addition, loss of connectivity may also reduce survival rates if individuals have to cross a hostile matrix, though this hypothesis has been rarely tested. Here, we address these issues by evaluating how individual survival and inter-patch movement probabilities of the European pond turtle (Emys orbicularis) are influenced by patch connectivity.
2. The study was carried out in a network of temporary ponds embedded in a heterogeneous agricultural matrix in southwestern Portugal, encompassing a period associated with a severe drought (2003–2005) and another with wetter climatic conditions (2010–2014). We mapped the location of ponds and land uses around each pond, and quantified connectivity among ponds using least-cost distances based on patch location and resistance to movement of different land uses. We then used multistate capture-recapture modelling to quantify how survival and movement of this freshwater turtle were related to different metrics of landscape connectivity, in the wet and dry periods.
3. We captured 221 pond turtles, including 89 juveniles, 58 females, and 74 males. Survival was higher in ponds more connected with other ponds, especially for juvenile turtles. The probability of movement between ponds decreased with increasing least-cost distances. Movement probabilities tended to be higher in the dry than in the wet period.
4. Our results support the idea that landscape connectivity affects both movement and survival rates in a patchy population inhabiting a temporary pond network. These effects are likely to be particularly marked in unstable freshwater systems like ours, where individuals may have to move widely to escape drying ponds during particularly dry years.
5. Overall, our findings suggest that focusing conservation efforts solely on protecting discrete freshwater habitats such as temporary ponds may be insufficient, requiring also due consideration of landscape connectivity offered by the surrounding agricultural matrix to assure long-term persistence of patchy populations inhabiting such habitats.

     

Patterns,origin and possible effects of sediment pollution in a Mediterranean lake

The influence of water permanence and high intra- and inter-annual hydrological variability on macrobenthos (organisms >1 mm) was studied using a taxonomical and a functional approach. The study was carried out in a Mediterranean salt marsh. Monthly samples of macrobenthic fauna were collected during two consecutive hydroperiods from six ponds with different water permanence (temporary, semi-permanent and permanent waters). Organisms were assigned to five functional response groups based on life-strategies according to their capacity to survive desiccation events, their dispersion capability and the necessity of water for their reproduction. Results from both approaches showed that the benthic community was more related to pond type than to intra- and inter-annual variability. The second aim was to analyse to which extent patterns in functional groups were determined by the existence of succession patterns or to environmental variability. In this sense, a clear succession pattern was not observed. In contrast, in most of the functional groups (4 out of 5), species within each functional group showed similar responses to water fluctuations. However, species of the fifth functional group, which comprised species without any particular adaptation to desiccation survival or avoidance, showed different responses to water level fluctuations. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

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.     

The temporal dynamics of temporary pond macroinvertebrate communities over a 10-year period

Ponds support a rich biodiversity. This arises in part because of the number and heterogeneity of ponds spatially throughout the landscape. Studies of ponds suggest that distinct communities develop within individual ponds but most examples are based on short-term 1- or 2-year surveys which cannot identify the effects of historic events upon contemporary communities. This study reports the development and turnover of the early summer macroinvertebrate communities in thirty small temporary ponds from their creation in 1994 over 10 years to 2004. Distinct pioneer communities established in the first year of the ponds’ creation, the first 3 years dominated by a fauna associated with long summer dry phases. Then a sustained period of inundation lasting 27 months from summer 1997–1999 resulted in establishment of many taxa associated with permanent ponds and loss of some temporary pond species. The re-establishment of summer dry phases in 1999 was associated with the loss of some but not all of the permanent water taxa and re-colonisation by some temporary water species creating new communities combining these different elements. The communities were not a linear successional sequence; the communities that re-assembled following resumption of dry phases reflected the contingent history of each pond and the effects of historic events. The longer term nature of the study showed that the characteristic heterogeneity of pond invertebrate communities occurs through time as well as spatially and that the richness and variety of contemporary communities, which is often hard to explain from snap-shot studies, is partly the result of historic events.