Predator–prey relationships within natural,restored, and created vernal pools

We performed a multiyear monitoring study to compare amphibian habitat quality among four natural, four restored, and six created pools. We used successful reproduction and metamorphosis of two vernal pool indicator species, the wood frog and spotted salamander, to represent desired outcomes. Ordination techniques were used to identify the aspects of habitat quality that were most correlated with desired outcomes. Previously published results indicated that pool depth, volume, and hydroperiod were among the best predictors of success, regardless of pool type. Observations in the first few years of monitoring also suggested that pools with longer hydroperiods had a greater abundance of aquatic predators of eggs and larvae of indicator species. This follow‐up study further explores and compares predator–prey relationships among pool types. We quantified within‐pool predator and prey abundance and diversity and collected another year of data on the reproductive success of indicator species. Our results confirmed that mean predator abundance was eight times higher in pools with longer hydroperiods. We documented a 96% decrease in wood frog survival rates in a semi‐permanent, natural pool following a 41% decrease in overhead canopy cover and an increase in green frog abundance. At the same time, wood frog reproductive success increased in nearby restored pools with lower predator abundance. Pools with the highest mean survival rates for the two indicator species combined were short‐ or long‐cycle pools (i.e. hydroperiod of 12–35 weeks) with low predator abundance (i.e. <1 organism L^?1) and greater proportions of arthropod prey relative to other food items.     

Hydrology,Physiochemistry, and Amphibians in Natural and Created Vernal Pool Wetlands

This study compared the hydrology, physiochemistry, and amphibian biomass between a complex of created vernal pools and a complex of natural vernal pools in 2007 in central Ohio, United States. Hydrologic connectivity of surface water and groundwater differed between the natural and the created pool complexes. Surface inundation duration for created pools exceeded that of natural pools, although spring water depths were similar. Dissolved oxygen (p= 0.05) and hourly temperature (p= 0.00) were 1.2% and 1.1% higher, respectively, in the created pools, and conductivity was 1.5% higher (p= 0.00) in the natural pools. Amphibian dip net results found no significant difference in biomass between natural and created pools or family (hylid, ranid, and ambystomatid) biomass in both pool types. Amphibian families were evenly represented by both capture methods in the created wetlands; however, the distribution of families was not even in natural pools and the proportion of ranids was four times greater for samples obtained by funnel traps than dip netting. Eleven years after construction, the created vernal pools did not mimic natural pools in surface inundation and groundwater–surface water exchange, dissolved oxygen, and water temperature. The created pools are perched wetlands and are never likely to mimic reference pool hydrology. Dissolved oxygen and temperature differences are likely due to the separation of surface water and groundwater in the created pools. However, the created pools exhibited a higher taxa diversity than the natural pools due to a more even distribution of organisms between the three families.     

Hydroperiods of created and natural vernal pools in central Ohio: A comparison of depth and duration of inundation

Water levels were recorded weekly from six natural vernal pools and 10 created vernal pools at two forested wetland complexes in central Ohio. Vernal pool median water depth and duration of inundation were significantly greater at the created vernal pools than at the natural vernal pools (α = 0.05, P < 0.05). The average period of inundation for created pools was 309 ± 32 days, compared with 250 ± 16 days for natural pools. The created pools produced a range of inundation times, from 163 to 365 days in length, with three pools permanently inundated.     

A Long‐Term Comparison of Hydrology and Plant Community Composition in Constructed Versus Naturally Occurring Vernal Pools

Successful restoration of ephemeral wetlands worldwide is particularly challenging, given the often‐precise relationship between hydrological features and plant community dynamics. Using a long‐term experiment in vernal pool restoration, we compare hydrological and vegetative characteristics of constructed pools with those of adjacent, naturally occurring reference pools. Although constructed and reference pools were similar in maximum water depth and duration of inundation at the beginning of our experiment in 2000, constructed pools were shallower and inundated for shorter periods by 2009. Native vernal pool species were able to establish populations in many constructed pools, and seeding sped their establishment. Comparing seeded plots in constructed pools with unseeded plots in reference pools, we found no significant difference in the cover of seeded species, native species, or exotic species in most years. In recent years, however, native species have declined in both constructed and reference pools. Finally, the cover of native vernal pool species was positively and non‐linearly associated with both water depth and seeding treatment. We conclude that the establishment of appropriate hydrological conditions was necessary, but not sufficient to promote successful performance of vernal pool species in constructed pools. Constructed pools with hydrologic conditions similar to those of reference pools were more likely to support populations of native vernal pool plant species, but only seeded pools were similar to reference pools in abundance of native cover. Most importantly, hydrological conditions in experimental pools have worsened since their construction, which may hamper persistence of native species in this restoration effort.     

The population biology of mitigation: impacts of habitat creation on an endangered plant species

Conservation practitioners widely agree that optimal conservation strategies will maximize the amount of genetic variation preserved in target taxa, but there is ongoing debate about how that variation should be distributed through restoration and mitigation activities. Here, we evaluate the impacts of ~10 years of mitigation on the population genetic structure of Limnanthes vinculans, a state- and federally-listed endangered plant species restricted to ephemeral vernal pool wetlands in the Santa Rosa Plain of California. Using microsatellite loci to estimate patterns of neutral molecular variation, we found that created pools support similar levels of variation in L. vinculans as natural pools. Habitat creation and seed translocation have not disrupted the largest-scale patterns of population structure across the species range, but a concentration of mitigation activity towards the range center has reduced the extent of isolation-by-distance operating in this region and shifted the location of at least one genetic boundary. Patterns of genetic variation among populations in remnant vernal pools reveal that gene flow has historically occurred beyond the scale of individual pools at the center of the species range, while small genetic populations have differentiated around the range margins. On average, L. vinculans in created pools exhibit less cover and more restricted local distributions than those in remnant pools, but these patterns were driven by two particularly productive natural sites rather than consistent differences between natural and created sites. We conclude that mitigation activities have changed the historical patterns of gene flow within the species range to a moderate degree, that these changes will likely impact remnant pools through gene flow, and that current created sites provide less heterogeneous habitat for L. vinculans than do natural pools. Studies that track individual plants will be needed to determine if the changes in gene flow due to mitigation will have positive or negative impacts on the demographic and microevolutionary trajectories of L. vinculans. More generally, this study provides a retrospective analysis of the outcome of managing an endangered plant species through intensive mitigation, and yields several insights to inform future conservation strategies.     

Livestock grazing affects vernal pool specialists more than habitat generalists in montane vernal pools

Questions

Do livestock grazing and seasonal precipitation structure species composition in montane vernal pools? Which grazing and precipitation variables best predict cover of vernal pool specialists and species with broader habitat requirements? Is vernal pool species diversity correlated with livestock exclosure, and at what spatial scales?

Location

Montane vernal pools, northeast California, USA.

Methods

Vegetation was sampled in 20 vernal pools, including pools where livestock had been excluded for up to 20 years We compared plant species composition, functional group composition and species diversity among sites that varied in grazing history and seasonal precipitation using CCA and LMM.

Results

Although vernal pool specialists were dominant in montane vernal pools, over a third of plant cover was comprised of species that occur over a broad range of wetland or upland environments. The species composition of vernal pool plant communities was influenced by both livestock grazing and precipitation patterns, however the relative effects of these environmental variables differed by functional group. Livestock exclosures favoured perennial vernal pool specialists over annual vernal pool specialists. In contrast, the cover of habitat generalists was more strongly influenced by seasonal precipitation than livestock grazing. At small spatial scales, species richness and diversity decreased as the number of years a pool had been fenced increased, but this relationship was not significant at a larger spatial scale.

Conclusions

Both livestock grazing and seasonal precipitation structure the montane vernal pool plant community. We found that livestock grazing promotes the cover of annual vernal pool specialists, but at the expense of perennial vernal pool specialists. Wetter vernal pools, however, support higher cover of wetland generalist species regardless of whether pools are grazed.     

Restoration genetics of the vernal pool endemic Lasthenia conjugens (Asteraceae)

Restoration of habitat for endangered species often involves translocation of seeds or individuals from source populations to an area targeted for revegetation. Long-term persistence of a species is dependent on the maintenance of sufficient genetic variation within and among populations. Thus, knowledge and maintenance of genetic variability within rare or endangered species is essential for developing effective conservation and restoration strategies. Genetic monitoring of both natural and restored populations can provide an assessment of restoration protocol success in establishing populations that maintain levels of genetic diversity similar to those in natural populations. California’s vernal pools are home to many endangered plants, thus conservation and restoration are large components of their management. Lasthenia conjugens (Asteraceae) is a federally endangered self-incompatible vernal pool annual with gravity- dispersed seeds. Using the molecular technique of intersimple sequence repeats (ISSRs), this study assessed levels and patterns of genetic variability present within natural and restored populations of L. conjugens. At Travis Air Force Base near Fairfield, California, a vernal pool restoration project is underway. Genetic success of the ecologically based seeding protocol was examined through genetic monitoring of natural and restored populations over a three-year period. Genetic diversity remained constant across the three sampled generations. Diversity was also widely distributed across all populations. We conclude that the protocol used to establish restored populations was successful in capturing similar levels and patterns of genetic diversity to those seen within natural pools. This study also demonstrates how genetic markers can be used to inform conservation and restoration decisions.     

Invasive plant feedbacks promote alternative states in California vernal pools

Invasive species have the potential to create positive feedbacks and push an ecosystem into an alternative state through a variety of mechanisms. Unless the drivers behind these feedbacks are understood, restoring a system to a more desirable state may not be possible. We used a long‐term vernal pool restoration project based out of Travis Airforce Base, Fairfield, CA, U.S.A. to examine natural pools dominated by either invasive or native plant communities, and restored pools predominately composed of invasive plants. We determined that plant community structure is drastically altered towards invasive grasses with the addition of a single centimeter of litter. In the absence of this litter layer, community structure was driven by a non‐native forb rather than native species. We also found that native plant‐dominated vernal pools have a longer inundation duration and are deeper compared to invasive‐dominated pools, regardless of construction status. These results suggest that once invasive grasses establish through lower inundation depths, their litter deposition can initiate a positive feedback to maintain an invasive alternative state. However, even after litter removal, non‐native forbs can replace the grasses causing a second alternative state still separate from the most desirable native dominated state. This study directly demonstrates that invasive species, and their positive feedbacks, may limit the success of ecological restoration. To effectively restore a system all constraints must be identified and removed before successful restoration can occur.     

Loss of biodiversity and hydrologic function in seasonal wetlands persists over 10 years of livestock grazing removal

Ecological restoration provides a means to increase biodiversity in ecosystems degraded by natural and human‐induced changes. In some systems, disturbances such as grazing can be key factors in the successful restoration of biodiversity and ecological function, but few studies have addressed this experimentally, especially over long time periods and at landscape scales. In this study, we excluded livestock grazing from plots within a grassland landscape containing vernal pools in the Central Valley of California for 10 years and compared vernal pool hydrology and plant community composition with areas grazed under an historic regime. In all 10 years, the relative cover of native plant species remained between 5 and 20% higher in the grazed versus ungrazed plots. This effect was particularly prominent on the pool edges, though evidence of invasion into the pool basins was evident later in the study. Native species richness was lower in the ungrazed plots with 10–20% fewer native species found in ungrazed versus grazed plots in all years except the first year of treatment. Ungrazed pools held water for a shorter period of time than pools grazed under an historic regime. By the ninth year of the study, ungrazed pools took up to 2 weeks longer to fill and dried down 1–2 weeks sooner at the end of the rainy season compared to grazed pools. The results of this study confirm that livestock grazing plays a key role in maintaining biodiversity and ecosystem function in vernal pools.     

Changes in vernal pool edaphic settings through mitigation at the project and landscape scale

Vernal pool mitigation is a highly controversial process that has been frequently criticized for its inability to adequately replicate the ecosystem functions of the original intact wetlands. We analyzed past mitigation practices in two rapidly growing counties in California's Great Central Valley to determine if mitigation procedures are re-arranging the vernal pool landscape by substituting more common or less ecologically significant pool types (as defined by soil type and geomorphology) for rarer or ecologically richer pool types. Results indicate that most development projects impacting vernal pools conduct at least a portion of their mitigation requirements at a site with similar edaphic settings. However, when examined at a landscape-scale across all development projects, the more common edaphic settings such as Northern Hardpan and Low Terrace pools are increasing while more rare types such as Northern Claypan and Volcanic Mudflow pools are decreasing. Results also show that Drainageway pools, a less-specialized pool type with generally lower species richness, are becoming more common through mitigation. These results are confirmed by an analysis of landscape diversity, which showed that overall landscape diversity was lower at mitigation sites than at project sites. Despite these results, the ecological significance of vernal pool mitigation practices remains unclear for several reasons. The lack of maps showing exact locations of vernal pools at project sites make it difficult to precisely determine vernal pool acreage and distribution among edaphic settings. Additionally, more research is needed to determine precise relationships between edaphic settings and species distributions and the effects of mitigation area management practices on species distribution and persistence.     

Multiscale resistant kernel surfaces derived from inferred gene flow: An application with vernal pool breeding salamanders

The importance of assessing spatial data at multiple scales when modelling species–environment relationships has been highlighted by several empirical studies. However, no landscape genetics studies have optimized landscape resistance surfaces by evaluating relevant spatial predictors at multiple spatial scales. Here, we model multiscale/layer landscape resistance surfaces to estimate resistance to inferred gene flow for two vernal pool breeding salamander species, spotted (Ambystoma maculatum) and marbled (A. opacum) salamanders. Multiscale resistance surface models outperformed spatial layers modelled at their original spatial scale. A resistance surface with forest land cover at a 500‐m Gaussian kernel bandwidth and normalized vegetation index at a 100‐m Gaussian kernel bandwidth was the top optimized resistance surface for A. maculatum, while a resistance surface with traffic rate and topographic curvature, both at a 500‐m Gaussian kernel bandwidth, was the top optimized resistance surface for A. opacum. Species‐specific resistant kernels were fit at all vernal pools in our study area with the optimized multiscale/layer resistance surface controlling kernel spread. Vernal pools were then evaluated and scored based on surrounding upland habitat (local score) and connectivity with other vernal pools on the landscape, with resistant kernels driving vernal pool connectivity scores. As expected, vernal pools that scored highest were in areas within forested habitats and with high vernal pool densities and low species‐specific landscape resistance. Our findings highlight the success of using a novel analytical approach in a multiscale framework with applications beyond vernal pool amphibian conservation.     

Effects of transgenic American chestnut leaf litter on growth and survival of wood frog larvae

Biotechnology offers a new approach for the restoration of tree species affected by exotic pathogens; however, nontarget impacts of this novel strategy on other organisms have not been comprehensively assessed. We evaluated the effect of transgenic American chestnut (Castanea dentata) leaf litter on the growth and survival of larval wood frogs (Lithobates sylvaticus), a forest‐dwelling amphibian species widely sympatric with American chestnut, that forage almost entirely on periphyton and litter detritus that accumulate in temporary vernal pools in forests. We reared wood frog larvae on Castanea leaf litter (American chestnut genetically engineered for blight tolerance, nontransgenic American chestnut, Chinese chestnut [Castanea mollissima], and an American–Chinese chestnut hybrid) and litter from two non‐Castanea, nontransgenic “control” tree species, coupled with two levels of supplementary food. We observed no differences in growth or survival of wood frog larvae reared on transgenic versus nontransgenic American chestnut leaves. Without supplementary food, wood frog larvae provided leaves from American chestnut (both types) developed faster and grew larger than those exposed to other leaf litter treatments. Results of this study provide preliminary evidence that (1) American chestnut may have formerly been an important source of food for forest‐dwelling amphibians and (2) transgenic American chestnut litter generated as part of chestnut restoration efforts is unlikely to present direct novel risks to developing amphibian larvae in the forest environment.     

Resilience of anostracan cysts to fire

California's Mediterranean ecosystems include shrubland and grassland vegetation types that are fire-prone. Dotted within this landscape are ephemeral wetlands called vernal pools. Since surrounding upland vegetation is adapted to survive fire, it is expected that vernal pool organisms should be able to survive as well. One group of animals common to vernal pools are anostracan crustaceans that survive the pool's dry period as encysted embryos. We hydrated anostracan cysts from the soil of a recently burned pool and from soil samples intentionally burned in a prescribed fire. We also sampled burned pools when refilled the next rainy season. We found that anostracan cysts in the soil can survive fire and that shrimp occur in pools in the first post-burn season. This information is important from a management perspective concerning fire effects, controlled or natural, on vernal pools and their rare and endangered species. This revised version was published online in August 2006 with corrections to the Cover Date.     

Protecting vernal pools: a model from Massachusetts,USA

Ephemeral ponded wetlands, often referred to as vernal pools in the wetland’s lexicon of the northeastern United States, are in need of protection from outright loss, catastrophic alteration, and disruption of natural processes in the surrounding landscape, because of their great importance to a characteristic wildlife fauna. The state of Massachusetts, USA provides a useful model for vernal pool protection. Vernal pools have been specifically protected under state wetlands regulations since 1987 and many of the state’s municipalities have added additional, more stringent protection under local wetland bylaws. Some vernal pools are also protected under federal wetlands protection law. The protection of vernal pools in Massachusetts is based on a certification process in which biological data are collected to demonstrate that a wetland provides vernal pool functions. Once certified, and if a pool meets jurisdictional requirements, regulatory restrictions are placed on development and other activities proposed within the pool. The regulatory protection for vernal pools in Massachusetts has resulted in the prevention of outright loss and degradation of many vernal pools across the state. However, a 2001 US Supreme Court decision, referred to as the SWANCC decision, may result in a loss of federal jurisdiction over many vernal pools, protection that is especially valuable for pools that do not meet state regulatory criteria. Of significant importance is a lack of protection in existing regulations of the upland non-breeding habitat surrounding vernal pools, which has been shown to be of critical importance to the animals that rely on the pools. Despite the shortcomings of the regulatory protection model in Massachusetts, it has been especially effective as a catalyst for environmental and wildlife awareness, public participation in the wetlands regulatory process, and as an extremely valuable tool in environmental education and outreach.     

Assessing Climate Change Impacts on Vernal Pool Ecosystems and Endemic Branchiopods

This study evaluated the hydrologic sensitivity of vernal pool ecosystems in the Central Valley of California to climatic changes projected for 2100. A vernal pool water-balance model was used to evaluate rain-fed vernal pools at four locations under future conditions projected by two contrasting global climate models. The potential for change in the duration of continuous inundation, frequency of reproductively suitable inundation events, and the seasonal distribution of inundation was quantified. The potential impact of hydrologic changes varied by species and by location. Three scales of response were identified: (a) At the regional scale, pools in the middle of the Central Valley near Merced were the most responsive to climatic changes. (b) At the local scale, smaller, shallower pools had the greatest potential to change the distribution of reproductively suitable habitat available to branchiopods. (c) At the individual pool scale, changes in precipitation will dominate changes in temperature, resulting in relatively linear responses in the duration of inundation. The ecological impact of these changes will be determined by a balance between the increasing suitability of vernal pools for branchiopod predators and the hydrologic improvement of currently marginal habitats.     

Success of Instream Habitat Structures After a 50-Year Flood in Gold Creek, Montana

Gold Creek, in western Montana, lost complexity and diversity of fish habitat following riparian logging activities, removal of instream wood, and subsequent scouring. In the 4.8-km study area, the stream was almost totally void of large woody debris (4.2 pieces/km) and associated pools (1.3 pools/km). We constructed 66 structures made of natural materials (rock and wood) that resulted in 61 new pools in the study area in an attempt to restore salmonid habitat in the fall of 1996. An estimated 50-year recurrence interval flood occurred in the following spring. Of the original 66 structures, 55 (85%) remained intact and stable. Laterally confined reaches retained significantly more pools than laterally extended reaches. Owing to a history of anthropogenic impacts in forested streams in the intermountain west, restoration efforts are needed. If instream structures are tailored to specific morphologic channel types, fish habitat restoration can be successful and withstand major floods.     

Behavioral, ecological, and molecular genetic analyses of reproductive strategies in the Amazonian dart-poison frog, Dendrobates ventrimaculatus

We report the first field and genetic studies of the reproductivestrategies of the Amazonian dart-poison frog Dendrobates ventrimaculatus,a species with biparental care. Neither males nor females arestrictly monogamous. Males are aggressively territorial, butsome females interact without aggression. Monitoring of breedingpools revealed high rates of multiple clutch deposition andhigh levels of larval cannibalism. Laboratory experiments confirmedlarval cannibalism and suggested a benefit to cannibals in increasedgrowth rate. Genetic analyses indicate that offspring from differentclutches in or above the same pool vary in relatedness and areon average intermediate in relatedness between individuals fromthe same clutch and unrelated individuals (from different pools).These data suggest that reproductive parasitism may be commonin this species     

Biogeochemical Hotspots in Forested Landscapes: The Role of Vernal Pools in Denitrification and Organic Matter Processing

Quantifying spatial and temporal heterogeneity in ecosystem processes presents a challenge for conserving ecosystem function across landscapes. In particular, many ecosystems contain small features that play larger roles in ecosystem processes than their size would indicate; thus, they may represent “hotspots” of activity relative to their surroundings. Biogeochemical hotspots are characterized as small features within a landscape that show comparatively high chemical reaction rates. In northeastern forests in North America, vernal pools are abundant, small features that typically fill in spring with snow melt and precipitation and dry by the end of summer. Ephemeral flooding alters soil moisture and the depth of the soil’s oxic/anoxic boundary, which may affect biogeochemical processes. We studied the effects of vernal pools on leaf-litter decomposition rates, soil enzyme activity, and denitrification in vernal pools to assess whether they function as biogeochemical hotspots. Our results indicate that seasonal inundation enhanced leaf-litter decomposition, denitrification, and enzyme activity in vernal pools relative to adjacent forest sites. Leaves in seasonally flooded areas decomposed faster than leaves in terra firme forest sites. Flooding also influenced the C, N, and P stoichiometry of decomposing leaf litter and explained the variance in microbial extracellular enzyme activity for phosphatase, β-d-glucosidase, and β-N-acetylglucosaminidase. Additionally, denitrification rates were enhanced by seasonal flooding across all of the study pools. Collectively, these data suggest that vernal pool ecosystems may function as hotspots of leaf-litter decomposition and denitrification and play a significant role in decomposition and nutrient dynamics relative to their size.