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Fruiting trees as dispersal foci in a semi-deciduous tropical forest.

Quantification of seed rain patterns is an initial step toward explaining variation in plant recruitment, and consequently, organization of forest communities. Spatially contagious patterns of seed deposition, where seeds are patchily dispersed with some sites receiving relatively high densities and others receiving low densities of seeds, may be a common phenomenon for which we have very little knowledge. For example, prior feeding events by frugivores (monkeys and birds) combined with transport and dispersal of seeds to other fruiting trees may result in the contagious deposition of non-conspecific seeds below them. Here, we examined whether fruiting trees act as dispersal foci in the semi-deciduous tropical rainforest of the Dja Reserve, Cameroon. Seed rain was sampled below the canopies of nine tree species: three typically dispersed by large, frugivorous birds, three dispersed by monkeys, and three dispersed by wind. We found no evidence that monkeys generate spatially contagious patterns of seed rain under fruiting trees at which they feed. However, we found that rates of deposition of non-conspecific seeds and species richness of seeds delivered by birds (hornbills and turacos) were significantly greater during fruiting than non-fruiting periods, and significantly greater under fruiting individuals of bird-dispersed tree species than under fruiting individuals of monkey- or wind-dispersed tree species. Additionally, during fruiting periods, the composition of non-conspecific seed rain under bird-dispersed tree species was more similar to other bird-dispersed trees than to monkey- or wind-dispersed tree species. The contagious dispersal of non-conspecific seeds to fruiting, bird-dispersed trees leads to higher seed densities under fruiting trees than those caused by local seed production. Non-conspecific seeds deposited in high densities may experience increased seed mortality even far from parent trees if predators are generalists. Alternatively, in the absence of complete density-dependent mortality, contagious seed dispersal could result in associations among species dispersed by the same dispersal agent.

Animals↗

Ecosystem implications of genetic variation in water-use of a dominant riparian tree.

Genetic variation in dominant species can affect plant and ecosystem functions in natural systems through multiple pathways. Our study focuses on how genetic variation in a dominant riparian tree ( Populus fremontii, P. angustifolia and their natural F(1) and backcross hybrids) affects whole-tree water use, and its potential ecosystem implications. Three major patterns were found. First, in a 12-year-old common garden with trees of known genetic makeup, hybrids had elevated daily integrated leaf-specific transpiration ( E(tl); P=0.013) and average canopy conductance ( G(c); P=0.037), with both E(tl) and G(c) approximately 30% higher in hybrid cross types than parental types. Second, delta(13)C values of leaves from these same trees were significantly more negative in hybrids ( P=0.004), and backcross hybrids had significantly more negative values than all other F(1) hybrid and parental types ( P<0.001). Third, in the wild, a similar pattern was found in leaf delta(13)C values where both hybrid cross types had the lowest values ( P<0.001) and backcross hybrids had lower delta(13)C values than any other tree type ( P<0.001). Our findings have two important implications: (1). the existence of a consistent genetic difference in whole-tree physiology suggests that whole-tree gas and water exchange could be another pathway through which genes could affect ecosystems; and (2). such studies are important because they seek to quantify the genetic variation that exists in basic physiological processes-such knowledge could ultimately place ecosystem studies within a genetic framework.

Carbon Isotopes↗

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities.

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen-Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen-Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary ( Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was approximately 1.5x farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen-Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen-Connell effects in Amazonian forests and may result in decreases in tree diversity.

Animals↗

Impacts of major predators on tropical agroforest arthropods: comparisons within and across taxa.

In food web studies, taxonomically unrelated predators are often grouped into trophic levels regardless of their relative importance on prey assemblages, multiple predator effects, or interactions such as omnivory. Ants and birds are important predators likely to differentially shape arthropod assemblages, but no studies have compared their effects on a shared prey base. In two separate studies, we excluded birds and ants from branches of a canopy tree ( Inga micheliana) in a coffee farm in Mexico for 2 months in the dry and wet seasons of 2002. We investigated changes in arthropod densities with and without predation pressure from (1) birds and (2) ant assemblages dominated by one of two ant species ( Azteca instabilis and Camponotus senex). We first analyzed individual effects of each predator (birds, Azteca instabilis, and C. senex) then used a per day effect metric to compare differences in effects across (birds vs ants) and within predator taxa (the two ant species). Individually, birds reduced densities of total and large arthropods and some arthropod orders (e.g., spiders, beetles, roaches) in both seasons. Azteca instabilis did not significantly affect arthropods (total, small, large or specific orders). Camponotus senex, however, tended to remove arthropods (total, small), especially in the dry season, and affected arthropod densities of some orders both positively and negatively. Predators greatly differed in their effects on Inga arthropods (for all, small, large, and individual orders of arthropods) both in sign (+/-) and magnitudes of effects. Birds had stronger negative effects on arthropods than ants and the two dominant ant species had stronger effects on arthropods in different seasons. Our results show that aggregating taxonomically related and unrelated predators into trophic levels without prior experimental data quantifying the sign and strengths of effects may lead to a misrepresentation of food web interactions.

Animals↗

Allocation to defense or growth in dipterocarp forest seedlings in Borneo.

We quantified the allocation of net production to plant secondary metabolites (especially condensed tannins and lignins) to evaluate the investment into defense mechanisms of three tropical forest canopy species in Borneo vis-à-vis the resource availability hypothesis. In particular, we focused on Borneo ironwood (Eusideroxylon zwageri, Lauraceae), which seemed to employ an extreme defensive strategy. The wood of this species is extremely durable and has a high specific gravity with a very slow growth rate. The allocation to defense by Borneo ironwood was compared to two emergent species of Dipterocarpaceae, the dominant family in this forest community. We conducted shade-house experiments on seedlings under four controlled conditions (two light levels x two nutrient levels) and showed that the growth rate of E. zwageri was much lower than those of the other two species, and it allocated more of its net production to leaves and roots than to stems. The concentrations of condensed tannins and lignins were very high in the leaves and stems of this species, at about 20 and 30%, respectively. In total, E. zwageri allocated a maximum of about 35% of its net production to defensive substances (i.e., condensed tannins + lignins). In contrast, the two dipterocarp species allocated about 20-25% of their net production to defensive substances. The condensed tannins in E. zwageri help to prolong the lifespan of the leaves, and the lignins in the stems enhance the durability of the wood. Thus, although E. zwageri grows very slowly, the allocation to defensive substances seems to be an advantageous strategy for survival under dark conditions.

Analysis of Variance↗

Cold and heat tolerance of drosophilid flies with reference to their latitudinal distributions.

The relation between thermal tolerance and latitudinal distribution was studied with 30 drosophilid species collected from the cool-temperate region (Sapporo), the warm-temperate region (Tokyo and Kyoto) and the subtropical region (Iriomote island) in Japan. In addition, intraspecific variation was examined for five species collected from two localities. The subtropical strains of Scaptodrosophila coracina, Drosophila bizonata and D. daruma were less tolerant to cold than their temperate strains. However, the difference of cold tolerance between these two geographic strains was much smaller than the difference between the species restricted to the subtropical region and those occurring in the temperate region. In D. auraria and D. suzukii, no difference was observed in thermal tolerance between their cool- and warm-temperate strains. Thus, geographic variation in thermal tolerance within species was low or negligible. Interspecific comparisons by phylogenetic independent contrasts revealed that species which had the northern boundaries of their distributions at higher latitudes were generally more tolerant to cold than those which had their boundaries at lower latitudes. However, the data for some species did not agree with this trend. The use of man-protected warm places for overwintering, competition or predation would also affect their distributions. It also appeared that species which had their southern boundaries at higher latitudes were generally more cold-tolerant. The acquisition of cold tolerance may lower a fly's capacity to compete, survive or reproduce in warmer climates. On the other hand, no relation was observed between heat tolerance and latitudinal distribution. Heat tolerance was higher in species inhabiting openlands or the forest canopy than in those inhabiting the forest understorey.

Adaptation, Physiological↗

Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution.

Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [ Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine ( Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2-4-mm diameter) collected at 20-30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance ( g(s)) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20-30 cm depth above -0.5 MPa in the old-growth Douglas-fir site and -1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about -2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g(s) decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.

Disasters↗

Evidence that fungal pathogens inhibit recruitment of a shade-intolerant tree, white birch ( Betula papyrifera), in understory habitats.

Evidence from tropical forests suggests understory habitats are associated with a high risk of disease, which may prevent the establishment of vulnerable tree species; in contrast, canopy gaps can act as refuges from these pathogens. However, few studies have investigated the impacts of pathogens on regeneration in temperate forests. To determine whether losses to fungi of seeds of Betula papyrifera, a light-loving species, varied between habitats that differed in their degree of openness, we applied fungicide to seeds buried in old fields, treefall gaps, and forest understory sites. We found that the application of fungicide significantly reduced losses in all habitats, relative to control values. This effect was habitat-dependent: the benefit of fungicide was greater in forest understory than in openings. This suggests that B. papyrifera is prevented from establishing in understory environments in part by its susceptibility to pathogen attack, and not solely because of a high light requirement.

Analysis of Variance↗

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests.

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.

Biodiversity↗

Microhabitat associations and seedling bank dynamics in a neotropical forest.

We conducted a rigorous test of tropical tree seedling microhabitat differentiation by examining microhabitat associations, survival and growth of established seedlings of ten tropical tree species representing a four-factor gradient in seed size. Eight microhabitat variables describing soil and light conditions were measured directly adjacent to each of 588 seedlings within twelve 10x100 m belt transects at Paracou, French Guiana, and at 264 reference points along the transects. From these measurements, we defined three principal components describing soil richness, soil softness and canopy openness. Six of ten species (in 9 of 30 total cases) were distributed non-randomly with respect to microhabitat along at least one principal component. However, few species demonstrated clear microhabitat specialization. All shifts in distribution relative to reference points were in the same direction (richer, softer soil). Furthermore, of 135 pairwise comparisons among the species, only 7 were significantly different. More than three-fourths of all seedlings (75.3%) survived over the 2-year monitoring period, but survival rates varied widely among species. In no case was the probability of survival influenced by any microhabitat parameter. Relative height growth rates for the seedlings over 2 years varied from -0.031 cm cm(-1) year(-1) (Dicorynia guianensis, Caesalpiniaceae) to 0.088 cm cm(-1) year(-1) (Virola michelii, Myristicaceae). In only 4 of 30 cases was height growth significantly associated with one of the three principal components. Because the conditions in this study were designed to maximize the chance of finding microhabitat differentiation among a group of species differing greatly in life history traits, the lack of microhabitat specialization it uncovered suggests that microhabitat partitioning among tropical tree species at the established seedling stage is unlikely to contribute greatly to coexistence among these species.

Ecosystem↗

Tight coupling between leaf area index and foliage N content in arctic plant communities.

The large spatial heterogeneity of arctic landscapes complicates efforts to quantify key processes of these ecosystems, for example productivity, at the landscape level. Robust relationships that help to simplify and explain observed patterns, are thus powerful tools for understanding and predicting vegetation distribution and dynamics. Here we present the same linear relationship between Leaf area index (LAI) and Total foliar nitrogen (TFN), the two factors determining the photosynthetic capacity of vegetation, across a wide range of tundra vegetation types in both northern Sweden and Alaska between leaf area indices of 0 and 1 m2 m(-2), which is essentially the entire range of leaf area index values for the Arctic as a whole. Surprisingly, this simple relationship arises as an emergent property at the plant community level, whereas at the species level a large variability in leaf traits exists. As the relationship between LAI and TFN exists among such varied ecosystems, the arctic environment must impose tight constraints on vegetation canopy development. This relationship simplifies the quantification of vegetation productivity of arctic vegetation types as the two most important drivers of productivity can be estimated reliably from remotely sensed NDVI images.

Alaska↗

Root hydraulic conductivity and whole-plant water balance in tropical saplings following a shade-to-sun transfer.

We hypothesized that pioneer and late successional species show different morphological and physiological responses in water use after gap formation. The magnitude of the responses was compared between two pioneer species (Macaranga gigantea and Trema orientalis) and four late successional species (Shorea sp.), in an experiment in which saplings were transferred from shade to sun. Although transpiration demand increased following the transfer, root hydraulic conductivity (Lp(r)) decreased. Lp(r) was sensitive to brief treatments with HgCl(2) (a specific inhibitor of aquaporins). This allows Lp(r) to be divided into two components: cell-to-cell and apoplastic pathways. The Lp(r) of cell-to-cell pathway decreased in all species following the transfer, relating to aquaporin depression in roots. Following the transfer, leaf osmotic potentials at full hydration decreased and both leaf mass per area [leaf mass/leaf area (LMA)] and fine-root surface area/leaf surface area (root SA/leaf SA) increased in almost all species, allowing saplings to compensate for the decrease in Lp(r). Physiologically, pioneer species showed larger decreases in Lp(r) and more effective osmotic adjustment than late successional species, and morphologically, pioneer species showed larger increases in root SA/leaf SA and LMA. Water balance at the whole-plant level should be regulated by coupled responses between the aboveground and the belowground parts. Interspecific differences in responses after gap formation suggest niche differentiation in water use between pioneer and late successional species in accordance with canopy-gap size.

Analysis of Variance↗

Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities.

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.

Analysis of Variance↗

Does predation contribute to tree diversity?

Seed and seedling predation may differentially affect competitively superior tree species to increase the relative recruitment success of poor competitors and contribute to the coexistence of tree species. We examined the effect of seed and seedling predation on the seedling recruitment of three tree species, Acer rubrum (red maple), Liriodendron tulipifera (yellow poplar), and Quercus rubra (northern red oak), over three years by manipulating seed and seedling exposure to predators under contrasting microsite conditions of shrub cover, leaf litter, and overstory canopy. Species rankings of seedling emergence were constant across microsites, regardless of exposure to seed predators, but varied across years. A. rubrum had the highest emergence probabilities across microsites in 1997, but Q. rubra had the highest emergence probabilities in 1999. Predators decreased seedling survival uniformly across species, but did not affect relative growth rates (RGRs). Q. rubra had the highest seedling survivorship across microsites, while L. tulipifera had the highest RGRs. Our results suggest that annual variability in recruitment success contributes more to seedling diversity than differential predation across microsites. We synthesized our results from separate seedling emergence and survival experiments to project seedling bank composition. With equal fecundity assumed across species, Q. rubra dominated the seedling bank, capturing 90% of the regeneration sites on average, followed by A. rubrum (8% of sites) and L. tulipifera (2% of sites). When seed abundance was weighted by species-specific fecundity, seedling bank composition was more diverse; L. tulipifera captured 62% of the regeneration sites, followed by A. rubrum (21% of sites) and Q. rubra (17% of sites). Tradeoffs between seedling performance and fecundity may promote the diversity of seedling regeneration by increasing the probability of inferior competitors capturing regeneration sites.

Ecosystem↗

Ant species confer different partner benefits on two neotropical myrmecophytes.

The dynamics of mutualistic interactions involving more than a single pair of species depend on the relative costs and benefits of interaction among alternative partners. The neotropical myrmecophytes Cordia nodosa and Duroia hirsuta associate with several species of obligately symbiotic ants. I compared the ant partners of Cordia and Duroia with respect to two benefits known to be important in ant-myrmecophyte interactions: protection against herbivores provided by ants, and protection against encroaching vegetation provided by ants. Azteca spp., Myrmelachista schumanni, and Allomerus octoarticulatus demerarae ants all provide the leaves of Cordia and Duroia some protection against herbivores. However, Azteca and Allomerus provide more protection than does Myrmelachista to the leaves of their host plants. Although Allomerus protects the leaves of its hosts, plants occupied by Allomerus suffer more attacks by herbivores to their stems than do plants occupied by other ants. Relative to Azteca or Allomerus, Myrmelachista ants provide better protection against encroaching vegetation, increasing canopy openness over their host plants. These differences in benefits among the ant partners of Cordia and Duroia are reflected in the effect of each ant species on host plant size, growth rate, and reproduction. The results of this study show how mutualistic ant partners can differ with respect to both the magnitude and type of benefits they provide to the same species of myrmecophytic host.

Analysis of Variance↗

Refuge function of marine algae complicates selection in an intertidal snail.

Species with restricted gene flow often show trait-shifts from one type of environment to another. In those rock-dwelling marine gastropods that lack larval dispersal, size generally decreases in wave-exposed habitats reducing risk of dislodgement, while increases in less exposed habitats to resist crab-crushing. In Littorina fabalis, however, snails of moderately exposed shores are generally much larger (11-14 mm) than snails of sheltered shores (5-8 mm). Observations from the White Sea (where crabs are not present) indicate that in the absence of crabs snails are small (6-7 mm) in both habitats. We assumed that the optimal size for L. fabalis in the absence of crabs is less than 8 mm, and thus that increased size in moderately exposed habitats in areas with crabs might be a response to crab predation. In a crab-rich area (Sweden) we showed that crab predation is an important mortality factor for this snail species in both sheltered and moderately exposed habitats. In sheltered habitats, snails were relatively more protected from crab-predation when dwelling on their habitual substrate, fucoid algae, than if experimentally tethered to rocks below the algae. This showed that algae function as snail refuges. Snail dislodgement increased, however, with wave exposure but tethering snails in moderately exposed habitats showed that large snails survived equally well on rocks under the algae as in the canopy of the algae. Thus in sheltered habitats a small snail size is favored, probably due to life-history reasons, while increased risk of being dislodged from the algae refuges promotes a large size in moderately exposed habitats. This study shows an example of selection of a trait depends on complex interactions of different factors (life-history optimization, crab predation, wave induced dislodgement and algal refuges).

Analysis of Variance↗

Ecophysiological responses to light availability in three Blechnum species (Pteridophyta, Blechnaceae) of different ecological breadth.

In Chilean evergreen temperate forest, fern species of the genus Blechnum occur in diverse microhabitats ranging from large gaps to heavily shaded understoreys. We hypothesised that differences in the ecological breadth of three co-occurring Blechnum species would be associated with differences in magnitude of ecophysiological responses to light availability. We quantified the field distribution of each species in relation to diffuse light availability (% canopy openness), and measured in situ variation in photosynthetic capacity (A), dark respiration (R (d)) and specific leaf area (SLA) across the light gradient. The response of SLA of each species was also evaluated in a common garden in two light conditions (understorey and forest edge). The three Blechnum species differed significantly in the range of light environments occupied (breadth: B. chilense > B. hastatum > B. mochaenum). Despite significant interspecific differences in average A and R (d), the response of these traits to light availability did not differ among species. However, there was significant interspecific variation in both the mean value and the plasticity of SLA to light availability, the species with least ecological breadth (B. mochaenum) showing a flatter reaction norm (lower response) than its two congeners. This pattern was also found in the common garden experiment. The adjustment of leaf morphology (SLA) to light availability appears to be an important mechanism of acclimation in these Blechnum species. The narrow range of light environments occupied by B. mochaenum may be at least partly attributable to its inability to display phenotypic plasticity in SLA to changes in light availability.

Adaptation, Physiological↗

Pulse additions of soil carbon and nitrogen affect soil nitrogen dynamics in an arid Colorado Plateau shrubland.

Biogeochemical cycles in arid and semi-arid ecosystems depend upon the ability of soil microbes to use pulses of resources. Brief periods of high activity generally occur after precipitation events that provide access to energy and nutrients (carbon and nitrogen) for soil organisms. To better understand pulse-driven dynamics of microbial soil nitrogen (N) cycling in an arid Colorado Plateau ecosystem, we simulated a pulsed addition of labile carbon (C) and N in the field under the canopies of the major plant species in plant interspaces. Soil microbial activity and N cycling responded positively to added C while NH (4) (+) -N additions resulted in an accumulation of soil NO (3) (-) . Increases in microbial activity were reflected in higher rates of respiration and N immobilization with C addition. When both C and N were added to soils, N losses via NH(3) volatilization decreased. There was no effect of soil C or N availability on microbial biomass N suggesting that the level of microbial activity (respiration) may be more important than population size (biomass) in controlling short-term dynamics of inorganic and labile organic N. The effects of C and N pulses on soil microbial function and pools of NH (4) (+) -N and labile organic N were observed to last only for the duration of the moisture pulse created by treatment addition, while the effect on the NO (3) (-) -N pool persisted after soils dried to pre-pulse moisture levels. We observed that increases in available C lead to greater ecosystem immobilization and retention of N in soil microbial biomass and also lowered rates of gaseous N loss. With the exception of trace gas N losses, the lack of interaction between available C and N on controlling N dynamics, and the subsequent reduction in plant available N with C addition has implications for the competitive relationships between plants species, plants and microbes, or both.

Bacterial Physiological Phenomena↗