Soil nitrogen availability in some arctic ecosystems in Northwest Alaska: responses to temperature and moisture
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Riverside terraces along the Kugururok River in the Noatak National Preserve provided an opportunity to study primary succession, considering general trends that apply across all terraces, and unique events that influence individual terraces. The 30-year-old willow/poplar (Salix spp., Populus balsamifera L.) terrace had no trees taller than 1.5 m; the abundant spruce trees were not tall enough to emerge from the canopy height of the willows and poplars, and moose (Alces alces [Clinton]) browsing limited the canopy height of these plants. The 75-year-old poplar/spruce (Picea glauca [Moench] Voss) terrace had a high density of poplars (> 1000/ha) and low density of spruce (125/ha); heavy browsing by moose reduced the density of poplar by about one-half. The removal of the poplar by moose in this stand resulted in sustained increases in growth of individual spruce trees. The 100-year-old younger spruce/poplar terrace had about twice as many spruce trees (1250/ha) as poplar trees (500/ha), and the spruce trees were larger on average than the poplar trees. In the 220+ year-old older spruce/poplar type, only a few poplars remained (about 25/ha), and the number of spruce trees (600/ha) was only half that of the younger stage, either from lower initial spruce density on this terrace, or increased mortality of spruce. The 240+ year-old spruce type was a second-generation forest, characterized by a high density (1950/ha) of small spruce trees, some of which were tilted, indicating discontinuous permafrost. Plant litterfall mass showed no strong trend with terrace age, although N content of litterfall appeared to decline by about 1/3 in the spruce-dominated stages. Fungal biomass increased with ecosystem age, whereas bacterial biomass and microfauna declined. We found no evidence of declining soil N supply in older stages, but fertilization experiments would be needed to determine if N limitation of productivity changed with ecosystem development. We conclude that the general successional trend of increased spruce dominance is robust for this location, but that unique events play important roles in determining tree densities and the timing of the shift in dominance from poplar to spruce. The arrival of moose in the 1970s accelerated dominance by spruce on young terraces.
Here we propose that an important cause of variation in species density may be prior environmental conditions that continue to influence current patterns. In this paper we investigated the degree to which species density varies with location within the landscape, independent of contemporaneous environmental conditions. The area studied was a coastal marsh landscape subject to periodic storm events. To evaluate the impact of historical effects, it was assumed that the landscape position of a plot relative to the river's mouth ('distance from sea') and to the edge of a stream channel ('distance from shore') would correlate with the impact of prior storm events, an assumption supported by previous studies. To evaluate the importance of spatial location on species density, data were collected from five sites located at increasing distances from the river's mouth along the Middle Pearl River in Louisiana. At each site, plots were established systematically along transects perpendicular to the shoreline. For each of the 175 Plots, we measured elevation, soil salinity, percent of plot recently disturbed, percent of sunlight captured by the plant canopy (as a measure of plant abundance), and plant species density. Structural equation analysis ascertained the degree to which landscape position variables explained variation in species density that could not be explained by current environmental indicators. Without considering landscape variables, 54% of the variation in species density could be explained by the effects of salinity, flooding, and plant abundance. When landscape variables were included, distance from shore was unimportant but distance from sea explained an additional 12% of the variance in species density (R2 of final model = 66%). Based on these results it appears that at least some of the otherwise unexplained variation in species density can be attributed to landscape position, and presumably previous storm events. We suggest that future studies may gain additional insight into the factors controlling current patterns of species density by examining the effects of position within the landscape.
Temperatures across the northern regions of North America have been increasing for 150 years, and forests have responded to this increase. In the Noatak National Preserve in Alaska, white spruce (Picea glauca [Moench] Voss) forests reach their northern limit, occurring primarily on well-drained sites and as gallery forests along streams. Rolling plateaus of tundra separate the white spruce forests into disjunct stands. We examined patterns of tree age, tree growth, and tree encroachment into tundra ecosystems in six stands along the Agashashok River. Warming over the past 150 years appears to have increased tree growth and resulted in forest expansion into adjacent tundra ecosystems. The forest/tundra ecotone shifted by about 80 to 100 m into the tundra in the past 200 years, as evidenced by declining maximum tree age with distance towards the tundra. The decadal-scale pattern of tree establishment at the farthest extent of trees into the tundra (the tundra-forest ecotone) correlated with the detrended growth index for trees within the forests; climate conditions that led to higher tree growth appeared to foster tree establishment in the tundra. This recent forest expansion has occurred across topographic boundaries, from well-drained soils on slopes onto poorly drained, flatter areas of tundra. Further expansion of the forests may be limited by more severe wind exposure and poor drainage that make the majority of tundra less suitable for trees.
At the northern limit of the boreal forest biome, alder ( Alnus crispa [Ait.] Pursh) shrubs occur in a variety of ecosystems. We assessed the effects of individual alder shrubs on soil properties and understory plant tissue nitrogen in floodplain terraces, valley slopes and tussock tundra ridges. The three ecosystems differed with respect to soil properties and abiotic conditions and supported distinct plant communities. Alder increased resin-exchangeable soil N and NO 3 production significantly in each ecosystem. The greatest difference between alder canopy and surrounding soil NO 3 measured both under field and laboratory conditions occurred in floodplain sites. The shrub effect on soil pH and soil organic matter was greatest on tundra ridges. Alder shrubs also influenced the nitrogen nutrition of plants growing beneath their canopies. Plants growing below alder canopies had higher foliar nitrogen concentration and natural abundance 15 N composition and lower carbon to nitrogen ratio than open-grown plants. Similar to soil N availability, understory plant leaf chemistry responded more to alder on floodplains than on slope or tundra ecosystems. This pattern suggests that understory plants rely more heavily on alder-fixed-N in this resource-poor ecosystem.
F oraging intensity and faecal inputs are important determinants of plant community response to herbivory. We used captive adult lesser snow geese ( Anser caerulescens caerulescens ), which feed on both above- and below-ground plant tissues, to manipulate foraging intensity and faecal inputs to plots in a sedge meadow in spring, 1996. We measured plant and soil characteristics throughout the growing season of 1996 and in August 1997. We analysed three contrasts: grazed plots versus ungrazed controls, plots with a short period of feeding (3 goose-hours) versus plots with a long period of feeding (6 goose-hours), and grazed plots with faeces versus grazed plots without faeces. Although grazed plots had an order of magnitude higher foraging intensity than that imposed by wild geese in the marsh, there was no effect of feeding on biomass or nitrogen concentration in the dominant species, Carex ramenskii and Triglochin maritimum , after one and two growing seasons. The amount of forage removed by geese did not differ between plots grazed for long and short periods, indicating that geese were able to remove little additional biomass after 3 hours due to low availability. Therefore, the amount of biomass removed by geese was a better indicator of foraging intensity than the time geese fed on plots. The presence of faeces had no effect on biomass or nitrogen concentration in Carex ramenskii or Triglochin maritimum , or on rates of net nitrogen mineralization in soils. Thus, faeces did not appear to increase nitrogen availability for plants in this marsh, probably because faecal density was low.
Consumptive and non-consumptive interactions of predators and prey can have strong direct and indirect effects on primary producers, such as stream algae. Increasing water temperatures may alter these interactions and thus influence productivity in streams. For each of 3 temperature treatments (‘ambient’, +2°C and +4°C), we measured the amount of algal biomass removed by grazing mayflies from 91 mesocosms after a 24-hour test period under 3 grazing treatments: lethal predators, non-lethal predators, and no predators. At all temperatures, grazers reduced algal biomass (p < 0.01), and the presence of lethal predators effectively dampened mayfly consumption of algae (p < 0.01). However, differences in algal biomass between lethal and non-lethal predator treatments were not significant, indicating that predators had no indirect behaviorally mediated effects on grazer consumption. Grazer removal of algal biomass marginally increased with increasing temperature (p = 0.051). We analyzed video data for changes in grazer foraging and drift behavior. Mayflies increased drift in the presence of lethal predators (p < 0.01) but not non-lethal predators, and no behavioral changes were seen with temperature increases. Mesocosms can help elucidate possible future shifts in trophic interactions due to climate warming. Yet, we found no evidence of indirect stonefly predator effects on grazing mayflies under these warming scenarios.
Joshua tree (Yucca brevifolia) is a distinctive and charismatic plant of the Mojave Desert. Although floral biology and seed production of Joshua tree and other yuccas are well understood, the fate of Joshua tree seeds has never been studied. We tested the hypothesis that Joshua tree seeds are dispersed by seed-caching rodents. We radioactively labelled Joshua tree seeds and followed their fates at five source plants in Potosi Wash, Clark County, Nevada, USA. Rodents made a mean of 30.6 caches, usually within 30 m of the base of source plants. Caches contained a mean of 5.2 seeds buried 3-30 nun deep. A variety of rodent species appears to have prepared the caches. Three of the 836 Joshua tree seeds (0.4%) cached germinated the following spring. Seed germination using rodent exclosures was nearly 15%. More than 82% of seeds in open plots were removed by granivores, and neither microsite nor supplemental water significantly affected germination. Joshua tree produces seeds in indehiscent pods or capsules, which rodents dismantle to harvest seeds. Because there is no other known means of seed dispersal, it is possible that the Joshua tree-rodent seed dispersal interaction is an obligate mutualism for the plant.
Agricultural development on floodplains contributes to hydrologic alteration and forest fragmentation, which may alter landscape-level processes. These changes may be related to shifts in the seed bank composition of floodplain wetlands. We examined the patterns of seed bank composition across a floodplain watershed by looking at the number of seeds germinating per m2 by species in 60 farmed and intact forested wetlands along the Cache River watershed in Illinois. The seed bank composition was compared above and below a water diversion (position), which artificially subdivides the watershed. Position of these wetlands represented the most variability of Axis I in a Nonmetric Multidimensional Scaling (NMS) analysis of site environmental variables and their relationship to seed bank composition (coefficient of determination for Axis 1: r2 = 0.376; Pearson correlation of position to Axis 1: r = 0.223). The 3 primary axes were also represented by other site environmental variables, including farming status (farmed or unfarmed), distance from the mouth of the river, latitude, and longitude. Spatial analysis based on Mantel correlograms showed that both water-dispersed and wind/water-dispersed seed assemblages had strong spatial structure in the upper Cache (above the water diversion), bur the spatial structure of water-dispersed seed assemblage was diminished in the lower Cache (below the water diversion), which lost floodpulsing. Bearing analysis also Suggested that water-dispersal process had a stronger influence on the overall spatial pattern of seed assemblage in the upper Cache, while wind/water-dispersal process had a stronger influence in the lower Cache. An analysis of the landscapes along the river showed that the mid-lower Cache (below the water diversion) had undergone greater land cover changes associated with agriculture than did the upper Cache watershed. Thus, the combination of forest fragmentation and hydrologic changes in the surrounding landscape may have had an influence on the seed bank composition and spatial distribution of the seed banks of the Cache River watershed. Our study suggests that the spatial pattern of seed bank composition may be influenced by landscape-level factors and processes.
Nitrogen enters terrestrial ecosystems through multiple pathways during primary succession. We measured accumulation of total soil nitrogen and changes in inorganic nitrogen (N) pools across a 300-y sequence of river terraces in northwest Alaska and assessed the contribution of the nitrogen-fixing shrub Shepherdia canadensis . Our work compared 5 stages of floodplain succession, progressing from a sparsely vegetated silt cap to dense shrubby vegetation, balsam poplar-dominated ( Populus balsamifera ) and white spruce-dominated ( Picea glauca ) mixed forests, and old-growth white spruce forest. Total soil N (0–30 cm depth) increased throughout the age sequence, initially by 2.4 g N·m −2 ·y −1 during the first 120 y of terrace development, then by 1.6 g N·m −2 ·y −1 during the subsequent 2 centuries. Labile soil N, measured by anaerobic incubation, increased most rapidly during the first 85 y of terrace formation, then remained relatively constant during further terrace development. On recently formed terraces, Shepherdia shrubs enriched soil N pools several-fold compared to soil beneath Salix spp. shrubs or intercanopy sites. Total and labile soil N accretion was proportional to Shepherdia cover during the first century of terrace development, and mineral soil δ 15 N content indicated that newly formed river terraces receive substantial N through N-fixation. About half the 600 g total N·m −2 accumulated across the river terrace chronosequence occurred during the 120 y when S. canadensis was dominant. Sediment deposited by periodic flooding continued to add N to terrace soils after the decline in Shepherdia abundance and may have contributed 25% of the total N found in the floodplain terrace soils.
Estimates of historical variability in river ecosystems are often lacking, but long-lived freshwater mussels could provide unique opportunities to understand past conditions in these environments. We applied dendrochronology techniques to quantify historical variability in growth-increment widths in valves (shells) of western pearlshell freshwater mussels ( Margaritifera falcata ). A total of 3 growth-increment chronologies, spanning 19 to 26 y in length, were developed. Growth was highly synchronous among individuals within each site, and to a lesser extent, chronologies were synchronous among sites. All 3 chronologies negatively related to instrumental records of stream discharge, while correlations with measures of water temperature were consistently positive but weaker. A reconstruction of stream discharge was performed using linear regressions based on a mussel growth chronology and the regional Palmer Drought Severity Index (PDSI). Models based on mussel growth and PDSI yielded similar coefficients of prediction (R 2 Pred ) of 0.73 and 0.77, respectively, for predicting out-of-sample observations. From an ecological perspective, we found that mussel chronologies provided a rich source of information for understanding climate impacts. Responses of mussels to changes in climate and stream ecosystems can be very site- and process-specific, underscoring the complex nature of biotic responses to climate change and the need to understand both regional and local processes in projecting climate impacts on freshwater species.
Attempts to understand how communities assemble following a disturbance are challenged by the difficulty of determining the relative importance of stochastic and deterministic processes. Biological legacies, which result from organisms that survive a disturbance, can favour deterministic processes in community assembly and improve predictions of successional trajectories. Recently disturbed ecosystems are often so rapidly colonized by propagules that the role of biological legacies is obscured. We studied biological legacies on a remote volcanic island in Alaska following a devastating eruption where the role of colonization from adjacent communities was minimized. The role of biological legacies in the near shore environment was not clear, because although some kelp survived, they were presumably overwhelmed by the many vagile propagules in a marine environment. The legacy concept was most applicable to terrestrial invertebrates and plants that survived in remnants of buried soil that were exposed by post-eruption erosion. If the legacy concept is extended to include ex situ survival by transient organisms, then it was also applicable to the island's thousands of seabirds, because the seabirds survived the eruption by leaving the island and have begun to return and rebuild their nests as local conditions improve. Our multi-trophic examination of biological legacies in a successional context suggests that the relative importance of biological legacies varies with the degree of destruction, the availability of colonizing propagules, the spatial and temporal scales under consideration, and species interactions. Understanding the role of biological legacies in community assembly following disturbances can help elucidate the relative importance of colonists versus survivors, the role of priority effects among the colonists, convergence versus divergence of successional trajectories, the influence of spatial heterogeneity, and the role of island biogeographical concepts.
The effects of site acidification on amphibian populations have been thoroughly addressed in the last decades. However, amphibians in naturally acidic environments, such as peatlands facing pressure from the peat mining industry, have received little attention. Through two field studies and an experiment, I assessed the use of bog habitats by the green frog (Rana clamitans melanota), a species sensitive to various forestry and peat mining disturbances. First, I compared the occurrence and breeding patterns of frogs in bog and upland ponds. I then evaluated frog movements between forest and bog habitats to determine whether they corresponded to breeding or postbreeding movements. Finally, I investigated, through a field experiment, the value of bogs as rehydrating areas for amphibians by offering living Sphagnum moss and two media associated with uplands (i.e., water with pH ca 6.5 and water-saturated soil) to acutely dehydrated frogs. Green frog reproduction at bog ponds was a rare event, and no net movements occurred between forest and bog habitats. However, acutely dehydrated frogs did not avoid Sphagnum. Results show that although green frogs rarely breed in bogs and do not move en masse between forest and bog habitats, they do not avoid bog substrates for rehydrating, despite their acidity. Thus, bogs offer viable summering habitat to amphibians, which highlights the value of these threatened environments in terrestrial amphibian ecology.