Invertebrate community composition differs between restored and natural vernal pools

The loss of freshwater wetlands worldwide has underscored the importance of restoration to enhance biodiversity and functional objectives. While aquatic invertebrate communities within restored perennial freshwaters are well studied, few studies have occurred in the greatly reduced habitat of seasonal wetlands, such as vernal pools. California vernal pools have experienced high habitat loss and support many threatened or endangered invertebrate species. We compared 90 natural and 90 restored vernal pools of different ages across 10 sites throughout California and Southern Oregon using the Sars' method. Large branchiopod abundance, total invertebrate abundance, class richness, and community composition were assessed between pool types (natural vs. restored) and along environmental gradients (e.g. site, pool depth, surface area, age since restoration). Large branchiopod and total invertebrate abundance were 215 and 274% higher in natural pools than restored pools, but class richness was not different. Community composition was significantly different and driven by greater abundances of vernal pool fairy shrimp, San Diego fairy shrimp, Ostracoda, Cladocera, and Copepoda in natural pools. Few environmental or habitat variables explained patterns in richness or abundance. Our work demonstrates that restored pools, even those decades old, are different than natural pools. Future mitigation and monitoring guidelines for restored vernal pools should include quantitative evaluations for aquatic invertebrates. Restored pools are not adequate compensation for lost natural pools because they do not have the same ecological functions and values.     

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.     

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.     

Cattle waste reduces plant diversity in vernal pool mesocosms

In California, much of the remaining vernal pool habitat is used for cattle grazing. Some studies suggest that grazing helps promote native plant diversity on grasslands, but the impact of grazing on plants that reside in pool basins is largely unknown. We investigated how one aspect of cattle grazing, the deposition of waste, affects these plant species by adding dung and urine to mesocosms lined with vernal pool soil. As a result of dung input, orthophosphate, conductivity, and turbidity increased in our mesocosms while dissolved oxygen decreased. Such changes in water quality are consistent with a shift toward a eutrophic state. Algal biomass and percent-cover also increased in dung-treated mesocosms. When the mesocosms dried, vascular plant species richness and percent-cover in dung-treated mesocosms were reduced by up to 54% and 87%, respectively. We attribute this to light attenuation by algal mats that flourished in the nutrient-enriched water. We also found that dung input caused significant, but weak, shifts in the composition of the vascular plant community. We conclude that cattle grazing may be detrimental to plant communities in vernal pools via increased nutrient loading, which promotes algal growth. Any beneficial effects of grazing may thus be limited to the surrounding grassland. Studies that examine the regional-scale impacts of grazing on vernal pool grasslands should separately consider the impacts to local-scale (i.e., within-pool) plant diversity, as most of the threatened and endangered plant species of California vernal pools reside primarily in pool basins.     

California vernal pool endemic responses to hydroperiod,plant thatch,and nutrients

Many endemic large branchiopods inhabit ephemeral freshwater ecosystems, including California vernal pools. Hydroperiod, inundation length, has been well studied in these systems that cycle between aquatic and terrestrial phases, but species’ responses to other ecological processes are still poorly known. For example, temporal (plant thatch from the previous terrestrial phase) and spatial (nutrient runoff) factors may have strong effects on emergence and population densities during the aquatic phase. We examined the effects of hydroperiod stability, thatch, and nutrients on the emergence and density of 4 vernal pool endemic species: Branchinecta lynchi (Anostraca), Linderiella occidentalis (Anostraca), Lepidurus packardi (Notostraca), and Cyzicus californicus (Spinicaudata). A full factorial mesocosm experiment was conducted which measured species densities, along with water quality variables. Hydroperiod and thatch differentially affected 3 of the 4 species based on emergence timing and life cycle. Treatments had effects on many water quality variables, and these variables were correlated with densities. These results highlight how hydroperiod stability along with other processes can affect large branchiopod species in temporary freshwater ecosystems. California vernal pools are a greatly reduced habitat rich in endemic and endangered species (including Branchinecta lynchi and Lepidurus packardi), and therefore, these results have implications for conservation and management.     

Spatial context strongly affects community composition of both passively and actively dispersing pool invertebrates in a highly heterogeneous landscape

相似文献   

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.     

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.     

Conservation of rare species with island-like distributions: A case study of Lasthenia conjugens (Asteraceae) using population genetic structure and the distribution of rare markers

Californian vernal pools, a patchy, island-like habitat, are endangered as a result of habitat destruction. Conservation of the remaining vernal pool habitat is essential for the persistence of several endangered species. We present the first study examining DNA-level genetic diversity within and among populations of a vernal pool plant species. We investigated genetic variation across eight populations of the US federally endangered vernal pool endemic Lasthenia conjugens (Asteraceae) using intersimple sequence repeat (ISSR) markers. Genetic diversity within the species was high (Nei's gene diversity estimate was 0.37), with moderate differentiation among populations (Bayesian F _ST analog of 0.124). Using an amova analysis, we found that the majority of the genetic variation (84%) was distributed within populations. There is a significant relationship between geographical distance and pairwise genetic differentiation as measured by the Bayesian estimate θ^B. The alternative hypotheses of historic geological processes within the Central Valley and contemporary gene flow are discussed as explanations of the data. Because of the vulnerability of the populations, we calculated a probability of loss for rare alleles (fragments) in the populations. Calculations show that sampling only one of the eight populations for ex-situ conservation or restoration will capture approximately 54% of the sampled rare fragments. We believe that one of the sampled populations has become extinct since it was sampled. When removing this population from the above-mentioned calculations, sampling one population will capture only 41.3% of the sampled rare fragments. We recommend sampling strategies for future conservation and restoration efforts of L. conjugens.     

Identification of co-occurring Branchinecta fairy shrimp species from encysted embryos using multiplex polymerase chain reaction

Morphological identification of many fairy shrimp species is difficult because distinguishing characters are restricted to adults. We developed two multiplex polymerase chain reaction assays that differentiate among three Branchinecta fairy shrimp with distributional overlap in southern California vernal pools. Two of the species are federally listed as threatened. Molecular identification of Branchinecta from cysts allows for species surveys to be conducted during the dry season, expanding the timeframe for population assessment and providing a less intrusive method of sampling sensitive vernal pool habitats.     

Comparing amphibian habitat quality and functional success among natural,restored, and created vernal pools

The fact that several vernal pool restoration and creation attempts in eastern Pennsylvania and New Jersey have been paired with conservation of natural pools in the same area provided a valuable research opportunity to compare amphibian habitat quality between project sites and natural reference pools. To measure desired outcomes, we used successful reproduction and metamorphosis of two vernal pool indicator species, the wood frog and spotted salamander. Although many previous studies indicate that restored and created pools rarely replace function lost in the destruction of natural pools, success of vernal pool indicator species was not necessarily related to pool type in this study. Results indicate a strong correlation between reproductive success for both species and vernal pool size (i.e. mean depth and volume), regardless of pool type. Although overall survival rates of wood frog larvae were significantly higher in natural pools with hydroperiods between 12 and 35 weeks, wood frogs were also successful in one restored and one created vernal pool. Salamander survival rates were highest in two natural and two created pools, which had in common both greater volumes and higher proportions of forest land cover in the surrounding 1,000 m. The documented success of vernal pool indicator species in two well‐established created pools demonstrates that pool creation can sometimes restore communities and ecological functions lost, especially when nearby natural pools are degraded or destroyed.     

Rain-pools on peat moorland as island habitats for midge larvae

A common aquatic habitat on peat moorland is a pool, often only a few centimetres deep. Many have a shorter average duration than that of the inhabitants but they are, nonetheless, inhabited by the larval stages of several chironomid species at moderately high densities. Because of their durational characteristics, these pools might be expected to have an endemic fauna, as is the case in similar ephemeral island habitats elsewhere. However, we find no such fauna. Indeed, we find no qualitative difference between them and permanent pools of the same substratum.We identify three reasons for the apparent absence of endemism. First, the midge fauna is able to survive dry periods in the moist mud, so that even the smallest pools are effectively permanent from the adaptive point of view. Second, although seasonal freezing eliminates the fauna of smaller pools, larvae are able to complete their development in the preceding summer. Third, nearby permanent pools which do not dry and in which the mud does not freeze, may act as refugia for re-invasion.     

Ecosystem-level effects of bioturbation by the tadpole shrimp <Emphasis Type="Italic">Lepidurus packardi</Emphasis> in temporary pond mesocosms

An example of ecosystem engineering gaining attention in aquatic systems is bioturbation, the disruption of sediment at the water–sediment interface due to burrowing and foraging. One consequence of bioturbation can be increased turbidity from suspended sediment, which generally inhibits macrophyte growth and reduces ecosystem functioning. Conversely, bioturbation may promote invertebrate species richness by unearthing dormant cysts. Temporary-pond crustaceans are not widely regarded as agents of bioturbation, but on the basis of aquaria observations we hypothesized that certain taxa can disturb the sediment and create highly turbid water. We tested this hypothesis by removing crustaceans from mesocosms lined with vernal pool soil. Compared to this treatment group, mesocosms containing crustaceans had extremely high turbidity from suspended sediment, as well as reduced total macrophyte cover. We also found clear compositional differences in macrophyte communities between treatments, driven largely by differences in water physicochemistry, including turbidity. Regression analysis linked most of the bioturbation to the endangered notostracan Lepidurus packardi Simon 1886, which was a strong predictor of turbidity in our mesocosms. We also found a trend toward increased crustacean species richness in our mesocosms in the presence of this taxon. An analysis of published data from King et al. (1996) suggests that this trend may extend to natural vernal pools. Overall, our results suggest that L. packardi may have large effects on vernal pool communities, likely mediated in part through its disturbing of the sediment.     

An historical interpretation of habitat use by frogs in a Central Amazonian Forest

Interpretations of habitat use in tropical frog assemblages have centred on resource partitioning and stressed the influence of interspecific interaction and climatic fluctuation on numbers of species using various habitats. We used audio strip transects and visual methods to determine the species composition, reproductive modes, and habitat occupancy patterns of the entire assemblage of frog species in 1900 hectares of primary forest north of Manaus in the central Amazon. We then compared taxon, reproductive mode, and habitat of species at six analogous lowland forest sites of similar species richness (five in the Amazon and one in Southeast Asia) to determine similarity of habitat use among sites and whether habitat is strongly associated with species» systematic positions. In all lowland Amazonian faunas, most species with aquatic development use pools, many species undergo some degree of terrestrial development, and few species are riparian or develop in streams. In contrast, about half the species in Southeast Asian assemblages are riparian and develop in streams, and few species develop terrestrially. Because reproductive mode and habitat associate strongly with taxon, patterns of habitat use observed at this regional scale are better explained by historical biogeography and differential rates of speciation than by proximal selection generated by contemporary environmental conditions. This study presents an inventory of frog species in a central Amazonian terre-firme forest and measurements of habitat availability and use by an entire assemblage of frogs throughout a large area (other portions of this study were published by Gascon, 1990, 1991; Zimmerman & Rodrigues, 1990; Zimmerman, 1991). We asked whether this local pattern of habitat occupancy differed from the regional Amazonian pattern and whether local species composition could be predicted from (sub)habitat composition. Viewing the assemblage at the local level did reveal species-(sub)habitat relationships masked at the broader regional level. About half the pool-breeders at the Manaus forest study sites would not use pools that could be flooded by a permanent stream; several species distinguished between permanent and temporary ponds; and some species occupied all available breeding habitat, whereas others occurred patchily. This pattern was maintained over four breeding seasons, and species composition could be predicted from (sub)habitat composition. Phylogeny was not a predictor of subhabitat occupancy. Perhaps species are phylogenetically constrained to develop in pool, stream, riparian, or terrestrial habitats, but contemporary selection governs their narrow distribution within these major habitat types. Finally, we asked whether anuran species richness in the central Amazon differs from that of the upper or lower Amazon. One genus, Eleutherodactylus , accounts for elevated species richness at upper Amazonian sites. Dry seasons in the central and lower Amazon are unlikely to restrict the spread of eleutherodactylines, which reproduce terrestrially. There are as many non-eleutherodactylines with terrestrial development at seasonal sites as there are at continually wet sites. Colonization history and the topography of central and lower Amazonia are more likely to limit eleutherodactyline richness.     

Assemblages of dytiscid predators and culicid prey in relation to environmental factors in natural and clear-cut boreal swamp forest pools

Assemblages of diving beetles (Dytiscidae) and mosquito immatures (Culicidae) were studied during 1987–1988 in 40 small, more or less temporary, snowmelt pools in spruce swamp forest and clearings at the Arctic Circle in Sweden. Larger pools were warmer than smaller ones, and clearing pools were warmer than forest pools. Temperature differences between pools remained high until late July. Twenty-one dytiscid species, representing three guilds, occurred in the pools, and individual pools had 1–13 species. Ten species occurred in both habitats. A few stenotopic species of boreal swamp forests had in clearing pools apparently been replaced by some species with a preference for more productive, often man-made habitats. Six Aedes species were collected in the clearing pools. Five of these were found in the forest pools, of which three had no mosquito larvae. Dytiscid assemblages in both habitats and culicid assemblages in clearing pools showed strong nested patterns.Abundance and species richness of both culicids and dytiscids were higher in clearing than in forest pools with an area >2 m² after that the effects of pool area had been accounted for (MANCOVA). In both habitats, abundance and species richness of both culicids and dytiscids were strongly and positively correlated with a linear combination of pool area, depth and temperature (Canonical Correlation). Increasing drought frequency of pools had a negative, less significant effect on the biota. In the clearing, the abiotic correlations with abundance were somewhat weakened chiefly by the relatively low abundance values from the largest pool.Distribution and mean abundance of individual dytiscid species were positively related in the clearing pools. Flying dytiscids were trapped in the larger (1.6 m²), but not in the smaller (0.07 m²) artificial pools, and the immigration rate was markedly higher on clearings than in forest. Dug pools were colonized faster on clearings than in forest. Even the flightless Hydroporus melanarius colonized dug pools during the first year.     

The effects of fire, local environment and time on ant assemblages in fens and forests

We investigated the effects of the abiotic environment, plant community composition and disturbance by fire on ant assemblages in two distinct habitat types in the Siskiyou Mountains in northern California and southern Oregon, USA. Sampling over 2 years in burned and unburned Darlingtonia fens and their adjacent upland forests, we found that the effects of disturbance by fire depended on habitat type. In forests, fire intensity predicted richness in ant assemblages in both years after the fire, and plant community composition predicted richness 2 years after the fire. No factors were associated with richness in the species‐poor fen ant assemblages. Species‐specific responses to both habitat type and disturbance by fire were idiosyncratic. Assemblage composition depended on habitat type, but not disturbance by fire, and the composition of each assemblage between years was more dissimilar in burned than unburned sites.     

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.