An important component of the hydrological niche involves the partitioning of water sources, but in landscapes characterized by shallow soils over fractured bedrock, root growth is highly constrained. We conducted a study to determine how physical constraints in the root zone affected the water use of three tree species that commonly coexist on the Edwards Plateau of central Texas; cedar elm (Ulmus crassifolia), live oak (Quercus fusiformis), and Ashe juniper (Juniperus ashei). The year of the study was unusually dry; minimum predawn water potentials measured in August were -8 MPa in juniper, less than -8 MPa in elm, and -5 MPa in oak. All year long, species used nearly identical water sources, based on stable isotope analysis of stem water. Sap flow velocities began to decline simultaneously in May, but the rate of decline was fastest for oak and slowest for juniper. Thus, species partitioned water by time when they could not partition water by source. Juniper lost 15-30 % of its stem hydraulic conductivity, while percent loss for oak was 70-75 %, and 90 % for elm. There was no tree mortality in the year of the study, but 2 years later, after an even more severe drought in 2011, we recorded 34, 14, 6, and 1 % mortality among oak, elm, juniper, and Texas persimmon (Diospyros texana), respectively. Among the study species, mortality rates ranked in the same order as the rate of sap flow decline in 2009. Among the angiosperms, mortality rates correlated with wood density, lending further support to the hypothesis that species with more cavitation-resistant xylem are more susceptible to catastrophic hydraulic failure under acute drought.
Seasonality in surface weather results in seasonal temperature and humidity changes in caves. Ecological and physiological differences among trogloxenes, troglophiles, and troglobionts result in species-dependent responses to this variability. To investigate these responses, we conducted five biological inventories in a marble cave in the Sierra Nevada Range, California, USA between May and December 2010. The cave was divided into six quadrats and temperature was continuously logged in each (humidity was logged at the entrance and in the deep cave). With increasing distance from the entrance, temperature changes were increasingly attenuated and lagged relative to surface temperature. Linear regressions were created to determine the relationship between measured environmental variables and diversity for cavernicoles (troglobionts and troglophiles) and trogloxenes cavewide and in the transition zone. Diversity for cavernicoles and trogloxenes peaked in the entrance and deep cave zones, respectively. Quadrat, date, 2-week antecedent temperature average, 2-week antecedent temperature range, and trogloxene abundance explained 76% of cavernicole diversity variability. Quadrat explained 55% of trogloxene diversity variability. In the transition zone, trogloxene abundance explained 26% of cavernicole variability and 2-week antecedent temperature and 2-week antecedent temperature range explained 40% of trogloxene variability. In the transition zone, trogloxene diversity was inversely related to 2-week antecedent temperature average and 2-week antecedent temperature range, suggesting that species were moving into the transition zone when temperature was most stable. In a CCA of cavernicoles distribution data and environmental variables, 35% of variation in species-specific distributions was attributable to quadrat, and non-significant percentages were explained by date and environmental variables. Differences in assemblage structure among quadrats were largely due to differences between distributions of trogloxenes and cavernicoles, but responses varied among species. Differences are likely due to ecological niche width, physiological constraints, and competition. cave climate; troglobiont; troglophile; Sequoia National Park, California, USA; community response; species interaction Tobin B.W., Hutchins B.T. and Schwartz B.F., 2013. Spatial and temporal changes in invertebrate assemblage structure from the entrance to deep-cave zone of a temperate marble cave.
The prevailing paradigm in subterranean ecology is that below-ground food webs are simple, limited to one or two trophic levels, and composed of generalist species because of spatio-temporally patchy food resources and pervasive energy limitation. This paradigm is based on relatively few studies of easily accessible, air-filled caves. However, in some subterranean ecosystems, chemolithoautotrophy can subsidize or replace surface-based allochthonous inputs of photosynthetically derived organic matter (OM) as a basal food resource and promote niche specialization and evolution of higher trophic levels. Consequently, the current subterranean trophic paradigm fails to account for variation in resources, trophic specialization, and food chain length in some subterranean ecosystems. We reevaluated the subterranean food web paradigm by examining spatial variation in the isotopic composition of basal food resources and consumers, food web structure, stygobiont species diversity, and chromophoric organic matter (CDOM), across a geochemical gradient in a large and complex groundwater system, the Edwards Aquifer in Central Texas (USA). Mean δ13C values of stygobiont communities become increasingly more negative along the gradient of photosynthetic OM sources near the aquifer recharge zone to chemolithoautotrophic OM sources closer to the freshwater-saline water interface (FWSWI) between oxygenated freshwater and anoxic, sulfide-rich saline water. Stygobiont community species richness declined with increasing distance from the FWSWI. Bayesian mixing models were used to estimate the relative importance of photosynthetic OM and chemolithoautorophic OM for stygobiont communities at three biogeochemically distinct sites. The contribution of chemolithoautotrophic OM to consumers at these sites ranged between 25% and 69% of total OM utilized and comprised as much as 88% of the diet for one species. In addition, the food web adjacent to the FWSWI had greater trophic diversity when compared to the other two sites. Our results suggest that diverse OM sources and in situ, chemolithoautotrophic OM production can support complex groundwater food webs and increase species richness. Chemolithoautotrophy has been fundamental for the long-term maintenance of species diversity, trophic complexity, and community stability in this subterranean ecosystem, especially during periods of decreased photosynthetic production and groundwater recharge that have occurred over geologic time scales.
Summary Sympatric species are expected to exhibit specialisation that reduces interspecific competition in environments with food resources that exhibit little spatial or temporal variation in availability, while sympatric species in more unpredictable environments should exhibit generalised feeding strategies to exploit a wide range of variable resources. We combined stable isotopes and quantitative mouthpart morphometric data to investigate trophic structure among sympatric species in a diverse subterranean amphipod assemblage in the Edwards Aquifer, Texas, U.S.A. Seven amphipod species occupied different regions of isotopic (δ13C and δ15N) space, suggesting the use of different food resources. Trophic position, measured as δ15N, was negatively correlated with planar area of the mandible and number of molar ridges, while it was positively correlated with incisor width. Reduced molar size and robust incisors are generally associated with predatory feeding strategies in non‐subterranean amphipods. δ13C exhibited non‐significant relationships with mouthpart morphology although one species, Texiweckeliopsis insolita, had significantly different mouthpart and δ13C values. Another species, Stygobromus russelli, had more generalised mouthparts, but isotope values indicate that it had a unique and unidentified feeding method and food source, illustrating the obfuscating effect of phylogeny on form–function relationships. Intraspecific relationships between body size and isotope values indicate that amphipod species showed little to moderate ontogenetic shifts in trophic position. Furthermore, body size did not predict trophic position when data were combined across species, suggesting that larger amphipods do not necessarily feed at higher trophic levels in this community. Our results indicate that sympatric subterranean amphipod species can exhibit specialised feeding strategies, suggesting that competition among species is driving niche partitioning. These results contradict the assumption that variable resource availability in groundwater habitats selects for trophic generalists among sympatric species.
Potential recharge through thick soils in mantled sinkholes was quantified using differential electrical resistivity tomography (ERT). Conversion of time series two-dimensional (2D) ERT profiles into 2D volumetric water content profiles using a numerically optimized form of Archie's law allowed us to monitor temporal changes in water content in soil profiles up to 9 m in depth. Combining Penman-Monteith daily potential evapotranspiration (PET) and daily precipitation data with potential recharge calculations for three sinkhole transects indicates that potential recharge occurred only during brief intervals over the study period and ranged from 19% to 31% of cumulative precipitation. Spatial analysis of ERT-derived water content showed that infiltration occurred both on sinkhole flanks and in sinkhole bottoms. Results also demonstrate that mantled sinkholes can act as regions of both rapid and slow recharge. Rapid recharge is likely the result of flow through macropores (such as root casts and thin gravel layers), while slow recharge is the result of unsaturated flow through fine-grained sediments. In addition to developing a new method for quantifying potential recharge at the field scale in unsaturated conditions, we show that mantled sinkholes are an important component of storage in a karst system.
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