Do geographically isolated wetlands influence landscape functions?

Geographically isolated wetlands (GIWs), those surrounded by uplands, exchange materials, energy, and organisms with other elements in hydrological and habitat networks, contributing to landscape functions, such as flow generation, nutrient and sediment retention, and biodiversity support. GIWs constitute most of the wetlands in many North American landscapes, provide a disproportionately large fraction of wetland edges where many functions are enhanced, and form complexes with other water bodies to create spatial and temporal heterogeneity in the timing, flow paths, and magnitude of network connectivity. These attributes signal a critical role for GIWs in sustaining a portfolio of landscape functions, but legal protections remain weak despite preferential loss from many landscapes. GIWs lack persistent surface water connections, but this condition does not imply the absence of hydrological, biogeochemical, and biological exchanges with nearby and downstream waters. Although hydrological and biogeochemical connectivity is often episodic or slow (e.g., via groundwater), hydrologic continuity and limited evaporative solute enrichment suggest both flow generation and solute and sediment retention. Similarly, whereas biological connectivity usually requires overland dispersal, numerous organisms, including many rare or threatened species, use both GIWs and downstream waters at different times or life stages, suggesting that GIWs are critical elements of landscape habitat mosaics. Indeed, weaker hydrologic connectivity with downstream waters and constrained biological connectivity with other landscape elements are precisely what enhances some GIW functions and enables others. Based on analysis of wetland geography and synthesis of wetland functions, we argue that sustaining landscape functions requires conserving the entire continuum of wetland connectivity, including GIWs.

A dynamic reference model: a framework for assessing biodiversity restoration goals in a fire-dependent ecosystem.

The use of reference models as templates of historical or natural conditions to assess restoration progress is inherently logical; however, difficulties occur in application because of the need to incorporate temporal variation in ecosystems caused by disturbance and succession, as well as seasonal, interannual, or decadal variability. The landscape-scale restoration of the globally threatened and fire-dependent longleaf pine ecosystem in the southeastern United States is an example in which restoration efforts are even more complicated by the limited availability of extant reference sites. This study uses the dynamic reference conceptual framework to assess the direction and rate of recovery with respect to biodiversity restoration goals using a 15-year vegetation data set from an experimental restoration treatment in fire-excluded, hardwood-encroached longleaf pine sandhills. We compared ground-cover vegetation response to midstory hardwood removal through herbicide application, mechanical removal, and fire only. Nonmetric multidimensional scaling ordinations and proportional similarity analyses suggest that, while vegetation changed in all treatments over time, no differences in species composition or hardwood density in the ground cover were attributable to hardwood reduction treatments after 15 years with frequent prescribed fire. Furthermore, the results of this study indicate that considerable variability is associated with reference sites over time. Sites identified in 1994 as attainable restoration targets had become a moving target themselves, changing in magnitude consistent with alterations in restoration plots attributable to treatment effects and shaped by the modest increase in fire frequency imposed since 1998. In a broad restoration context, this study demonstrates a conceptual framework to better understand and integrate the range of spatial and temporal variation associated with the best available reference sites. It also illustrates a practical tool for statistically defining reference sites and for measuring restoration success in continually changing conditions that should be widely applicable to other ecosystems and restoration goals.

Relative effects of disturbance on red imported fire ants and native ant species in a longleaf pine ecosystem.

The degree to which changes in community composition mediate the probability of colonization and spread of non-native species is not well understood, especially in animal communities. High species richness may hinder the establishment of non-native species. Distinguishing between this scenario and cases in which non-native species become established in intact (lacking extensive anthropogenic soil disturbance) communities and subsequently diminish the abundance and richness of native species is challenging on the basis of observation alone. The red imported fire ant (Solenopsis invicta), an invasive species that occurs throughout much of the southeastern United States, is such an example. Rather than competitively displacing native species, fire ants may become established only in disturbed areas in which native species richness and abundance are already reduced. We used insecticide to reduce the abundance of native ants and fire ants in four experimental plots. We then observed the reassembly and reestablishment of the ants in these plots for 1 year after treatment. The abundance of fire ants in treated plots did not differ from abundance in control plots 1 year after treatment. Likewise, the abundance of native ants increased to levels comparable to those in control plots after 1 year. Our findings suggest that factors other than large reductions in ant abundance and species density (number of species per unit area) may affect the establishment of fire ants and that the response of native ants and fire ants to disturbance can be comparable.

Effects of resource availability on seedling recruitment in a fire-maintained savanna.

The herbaceous ground cover of the longleaf pine ecosystem harbors the highest plant species richness in North America, with up to 50 species per square meter, but the mechanisms that regulate this diversity are not well understood. In this system, variability in seedling recruitment events may best explain the extremely high small-scale species richness and its relationship to soil moisture and system net primary productivity. To understand the potential mechanistic controls on species richness, we used a long-term resource manipulation study across a natural soil moisture gradient to assess environmental controls on seedling recruitment. We considered the availability of resources to be an indicator of seedling safe-site supply, and also manipulated seed availability to examine the relative importance of recruitment limitations on seedling diversity. We found that water availability regulated the number of species in the seedling community regardless of the underlying natural moisture gradient, and that this effect may result from differential responses of seedling guilds to resource availability. Water supply was more important than seed supply in determining seedling establishment, suggesting that appropriate sites for regeneration are a factor limiting seedling success. This is the first study that shows that the episodic supply of microsites for recruitment could influence species richness in the highly threatened and biodiverse longleaf pine savanna.

Enhancing protection for vulnerable waters.

Governments worldwide do not adequately protect their limited freshwater systems and therefore place freshwater functions and attendant ecosystem services at risk. The best available scientific evidence compels enhanced protections for freshwater systems, especially for impermanent streams and wetlands outside of floodplains that are particularly vulnerable to alteration or destruction. New approaches to freshwater sustainability - implemented through scientifically informed adaptive management - are required to protect freshwater systems through periods of changing societal needs. One such approach introduced in the US in 2015 is the Clean Water Rule, which clarified the jurisdictional scope for federally protected waters. However, within hours of its implementation litigants convinced the US Court of Appeals for the Sixth Circuit to stay the rule, and the subsequently elected administration has now placed it under review for potential revision or rescission. Regardless of its outcome at the federal level, policy and management discussions initiated by the propagation of this rare rulemaking event have potential far-reaching implications at all levels of government across the US and worldwide. At this timely juncture, we provide a scientific rationale and three policy options for all levels of government to meaningfully enhance protection of these vulnerable waters. A fourth option, a 'do-nothing' approach, is wholly inconsistent with the well-established scientific evidence of the importance of these vulnerable waters.

Experimental test for facilitation of seedling recruitment by the dominant bunchgrass in a fire-maintained savanna.

Facilitative interactions between neighboring plants can influence community composition, especially in locations where environmental stress is a factor limiting competitive effects. The longleaf pine savanna of the southeastern United States is a threatened and diverse system where seedling recruitment success and understory species richness levels are regulated by the availability of moist microsites. We hypothesized that the dominant bunch grass species (Aristida stricta Michx.) would facilitate moist seedling microsites through shading, but that the effect would depend on stress gradients. Here, we examined the environmental properties modified by the presence of wiregrass and tested the importance of increased shade as a potential facilitative mechanism promoting seedling recruitment across spatial and temporal stress gradients. We showed that environmental gradients, season, and experimental water manipulation influence seedling success. Environmental properties were modified by wiregrass proximity in a manner that could facilitate seedling success, but we showed that shade alone does not provide a facilitative benefit to seedlings in this system.

Growth and photosynthetic responses of the federally endangered shrub, Lindera melissifolia (Lauraceae), to varied light environments.

Photosynthetic, morphological, and growth responses to light environment can be useful measurements to determine favorable habitat conditions for the conservation of endangered species. For Lindera melissifolia (Walt.) Blume, we compared morphological and photosynthetic responses under natural and controlled light regimes, and growth under three light treatments: 100%, 42%, and 19% full sunlight. Typical sun-shade morphological responses to decreasing light levels included decreased stomatal density, increased specific leaf area, and increased leaf area ratio. Photosynthetic capacity (3-6 µmol CO(2) · m(-2) · s(-1)) was consistent with other shade-tolerant species. Light-saturated rates of photosynthesis of experimental plants increased with increasing light up to 42% sunlight, but declined at 100% sunlight. The 100% light treatment also resulted in lower plant biomass, primarily from a reduction in root biomass. Results indicate that canopy conditions at levels below 40% sunlight are optimal for plant growth and should be considered in management and reintroduction efforts for this species.