Showing papers in "Oecologia in 2010"

A consistent terminology for quantifying species diversity? Yes, it does exist

Abstract: The prevailing terminological confusion around the concept ‘diversity’ has hampered accurate communication and caused diversity issues to appear unnecessarily complicated. In fact, a consistent terminology for phenomena related to (species) diversity is already available. When this terminology is adhered to, diversity emerges as an easily understood concept. It is important to differentiate between diversity itself and a diversity index: an index of something is just a surrogate for the thing itself. The conceptual problem of defining diversity also has to be separated from the practical problem of deciding how to adequately quantify diversity for a community of interest. In practice, diversity can be quantified for any dataset where units of observation (such as individuals) have been classified into types (such as species). All that needs to be known is what proportion of the observed units belong to a type of mean abundance. Diversity equals the inverse of this mean, and it quantifies the effective number of the types of interest. In ecology, interest often (but not always) focuses on species diversity. If the dataset consists of (or gets divided into) subunits, then the total effective number of species (gamma diversity) can be partitioned into the effective number of compositionally distinct subunits (beta diversity) and the mean effective number of species per such subunit (alpha diversity). Species richness is related to species diversity, but they are not the same thing; richness does not take the proportional abundances into account and is therefore the actual—rather than the effective—number of types. Most of the phenomena that have been called ‘beta diversity’ in the past do not quantify an effective number of types, so they should be referred to by names other than ‘diversity’ (for example, species turnover or differentiation).

Geographic and temporal correlations of mammalian size reconsidered: a resource rule

Abstract: The tendency of mammals to increase or decrease body size with respect to geography or time depends on the abundance, availability, and size of resources This dependency accounts for a change in mass with respect to geography, including latitude (Bergmann's rule), a desert existence, and life on oceanic islands (the island rule), as well as in a seasonal anticipation of winter (Dehnel's phenomenon) and a tendency for some lineages to increase in mass through time (Cope's rule) Such a generalized pattern could be called the "resource rule," reflecting the controlling effect of resource availability on body mass and energy expenditure The correlation of mammalian size with geography and time reflects the impact of temperature, rainfall, and season on primary production, as well as the necessity in the case of some species to share resources with competitors The inability of the constituent "rules" to account for all size trends often results from unique patterns of resource availability

Fine root decomposition rates do not mirror those of leaf litter among temperate tree species

Abstract: Elucidating the function of and patterns among plant traits above ground has been a major research focus, while the patterns and functioning of belowground traits remain less well understood. Even less well known is whether species differences in leaf traits and their associated biogeochemical effects are mirrored by differences in root traits and their effects. We studied fine root decomposition and N dynamics in a common garden study of 11 temperate European and North American tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pseudotsuga menziesii, Quercus robur, Quercus rubra and Tilia cordata) to determine whether leaf litter and fine root decomposition rates are correlated across species as well as which species traits influence microbial decomposition above versus below ground. Decomposition and N immobilization rates of fine roots were unrelated to those of leaf litter across species. The lack of correspondence of above- and belowground processes arose partly because the tissue traits that influenced decomposition and detritus N dynamics different for roots versus leaves, and partly because influential traits were unrelated between roots and leaves across species. For example, while high hemicellulose concentrations and thinner roots were associated with more rapid decomposition below ground, low lignin and high Ca concentrations were associated with rapid aboveground leaf decomposition. Our study suggests that among these temperate trees, species effects on C and N dynamics in decomposing fine roots and leaf litter may not reinforce each other. Thus, species differences in rates of microbially mediated decomposition may not be as large as they would be if above- and belowground processes were working in similar directions (i.e., if faster decomposition above ground corresponded to faster decomposition below ground). Our results imply that studies that focus solely on aboveground traits may obscure some of the important mechanisms by which plant species influence ecosystem processes.

The role of habitat quality in fragmented landscapes: a conceptual overview and prospectus for future research.

Abstract: There is increasing empirical evidence that the quality of habitat patches (determined by either habitat degradation or natural heterogeneity in the quality of habitat) plays an important role in determining species distribution patterns and in regulating spatial dynamics in fragmented landscapes. However, to date, most of the debate has focused on whether or not to include habitat variables in fragmentation studies, and we still lack general conclusions as well as standard and robust research approaches. In this paper we show how a weak conceptualization of “patch quality” and the inappropriate choice of target surrogate variables (e.g., density is often used as an indicator of patch quality) have mainly produced case-specific results, rather than general conclusions. We then identify weaknesses in the inclusion of habitat quality measurements within fragmentation studies. In particular, we focus on: (1) the lack of appropriate experimental design, outlining how few studies have actually included a gradient of habitat quality in their sample; (2) the lack of fundamental information provided (e.g., lack of standard outputs), which in turn hampers the possibility of carrying out meta-analyses. We finally synthesize available knowledge from empirical studies and highlight the different conceptual frameworks needed for patch occupancy versus patch use studies.

Leaf litter decomposition in temperate deciduous forest stands with a decreasing fraction of beech (Fagus sylvatica)

Abstract: We hypothesised that the decomposition rates of leaf litter will increase along a gradient of decreasing fraction of the European beech (Fagus sylvatica) and increasing tree species diversity in the generally beech-dominated Central European temperate deciduous forests due to an increase in litter quality. We studied the decomposition of leaf litter including its lignin fraction in monospecific (pure beech) stands and in stands with up to five tree genera (Acer spp., Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Tilia spp.) using a litterbag approach. Litter and lignin decomposition was more rapid in stand-representative litter from multispecific stands than in litter from pure beech stands. Except for beech litter, the decomposition rates of species-specific tree litter did not differ significantly among the stand types, but were most rapid in Fraxinus excelsior and slowest in beech in an interspecific comparison. Pairwise comparisons of the decomposition of beech litter with litter of the other tree species (except for Acerplatanoides) revealed a “home field advantage” of up to 20% (more rapid litter decomposition in stands with a high fraction of its own species than in stands with a different tree species composition). Decomposition of stand-representative litter mixtures displayed additive characteristics, not significantly more rapid than predicted by the decomposition of litter from the individual tree species. Leaf litter decomposition rates were positively correlated with the initial N and Ca concentrations of the litter, and negatively with the initial C:N, C:P and lignin:N ratios. The results support our hypothesis that the overall decomposition rates are mainly influenced by the chemical composition of the individual litter species. Thus, the fraction of individual tree species in the species composition seems to be more important for the litter decomposition rates than tree species diversity itself.

The effects of top-down versus bottom-up control on benthic coral reef community structure.

Abstract: While climate change and associated increases in sea surface temperature and ocean acidification, are among the most important global stressors to coral reefs, overfishing and nutrient pollution are among the most significant local threats. Here we examined the independent and interactive effects of reduced grazing pressure and nutrient enrichment using settlement tiles on a coral-dominated reef via long-term manipulative experimentation. We found that unique assemblages developed in each treatment combination confirming that both nutrients and herbivores are important drivers of reef community structure. When herbivores were removed, fleshy algae dominated, while crustose coralline algae (CCA) and coral were more abundant when herbivores were present. The effects of fertilization varied depending on herbivore treatment; without herbivores fleshy algae increased in abundance and with herbivores, CCA increased. Coral recruits only persisted in treatments exposed to grazers. Herbivore removal resulted in rapid changes in community structure while there was a lag in response to fertilization. Lastly, re-exposure of communities to natural herbivore populations caused reversals in benthic community trajectories but the effects of fertilization remained for at least 2 months. These results suggest that increasing herbivore populations on degraded reefs may be an effective strategy for restoring ecosystem structure and function and in reversing coral-algal phase-shifts but that this strategy may be most effective in the absence of other confounding disturbances such as nutrient pollution.

Altitudinal patterns of tick and host abundance: a potential role for climate change in regulating tick-borne diseases?

Abstract: The impact of climate change on vector-borne infectious diseases is currently controversial. In Europe the primary arthropod vectors of zoonotic diseases are ticks, which transmit Borrelia burgdorferi sensu lato (the agent of Lyme disease), tick-borne encephalitis virus and louping ill virus between humans, livestock and wildlife. Ixodes ricinus ticks and reported tick-borne disease cases are currently increasing in the UK. Theories for this include climate change and increasing host abundance. This study aimed to test how I. ricinus tick abundance might be influenced by climate change in Scotland by using altitudinal gradients as a proxy, while also taking into account the effects of hosts, vegetation and weather effects. It was predicted that tick abundance would be higher at lower altitudes (i.e. warmer climates) and increase with host abundance. Surveys were conducted on nine hills in Scotland, all of open moorland habitat. Tick abundance was positively associated with deer abundance, but even after taking this into account, there was a strong negative association of ticks with altitude. This was probably a real climate effect, with temperature (and humidity, i.e. saturation deficit) most likely playing an important role. It could be inferred that ticks may become more abundant at higher altitudes in response to climate warming. This has potential implications for pathogen prevalence such as louping ill virus if tick numbers increase at elevations where competent transmission hosts (red grouse Lagopus lagopus scoticus and mountain hares Lepus timidus) occur in higher numbers.

Isotopic values of plants in relation to water availability in the Eastern Mediterranean region

Abstract: Plant C and N isotope values often correlate with rainfall on global and regional scales. This study examines the relationship between plant isotopic values and rainfall in the Eastern Mediterranean region. The results indicate significant correlations between both C and N isotope values and rainfall in C3 plant communities. This significant relationship is maintained when plant communities are divided by plant life forms. Furthermore, a seasonal increase in C isotope values is observed during the dry season while N isotope values remain stable across the wet and dry seasons. Finally, the isotopic pattern in plants originating from desert environments differs from those from Mediterranean environments because some desert plants obtain most of their water from secondary sources, namely water channeled by local topographic features rather than direct rainfall. From these results it can be concluded that water availability is the primary factor controlling C and N isotope variability in plant communities in the Eastern Mediterranean.

Consumption of a nectar alkaloid reduces pathogen load in bumble bees.

Abstract: Diet has a significant effect on pathogen infections in animals and the consumption of secondary metabolites can either enhance or mitigate infection intensity. Secondary metabolites, which are commonly associated with herbivore defense, are also frequently found in floral nectar. One hypothesized function of this so-called toxic nectar is that it has antimicrobial properties, which may benefit insect pollinators by reducing the intensity of pathogen infections. We tested whether gelsemine, a nectar alkaloid of the bee-pollinated plant Gelsemium sempervirens, could reduce pathogen loads in bumble bees infected with the gut protozoan Crithidia bombi. In our first laboratory experiment, artificially infected bees consumed a daily diet of gelsemine post-infection to simulate continuous ingestion of alkaloid-rich nectar. In the second experiment, bees were inoculated with C. bombi cells that were pre-exposed to gelsemine, simulating the direct effects of nectar alkaloids on pathogen cells that are transmitted at flowers. Gelsemine significantly reduced the fecal intensity of C. bombi 7 days after infection when it was consumed continuously by infected bees, whereas direct exposure of the pathogen to gelsemine showed a non-significant trend toward reduced infection. Lighter pathogen loads may relieve bees from the behavioral impairments associated with the infection, thereby improving their foraging efficiency. If the collection of nectar secondary metabolites by pollinators is done as a means of self-medication, pollinators may selectively maintain secondary metabolites in the nectar of plants in natural populations.

The empirical Bayes approach as a tool to identify non-random species associations

Abstract: A statistical challenge in community ecology is to identify segregated and aggregated pairs of species from a binary presence-absence matrix, which often contains hundreds or thousands of such potential pairs. A similar challenge is found in genomics and proteomics, where the expression of thousands of genes in microarrays must be statistically analyzed. Here we adapt the empirical Bayes method to identify statistically significant species pairs in a binary presence-absence matrix. We evaluated the per- formance of a simple confidence interval, a sequential Bonferroni test, and two tests based on the mean and the confidence interval of an empirical Bayes method. Observed patterns were compared to patterns generated from null model randomizations that preserved matrix row and column totals. We evaluated these four methods with random matrices and also with random matrices that had been seeded with an additional segregated or aggregated species pair. The Bayes methods and Bonferroni correc- tions reduced the frequency of false-positive tests (type I error) in random matrices, but did not always correctly identify the non-random pair in a seeded matrix (type II error). All of the methods were vulnerable to identifying spurious secondary associations in the seeded matrices. When applied to a set of 272 published presence-absence matrices, even the most conservative tests indicated a fourfold increase in the frequency of perfectly segregated ''checkerboard'' species pairs compared to the null expectation, and a greater predominance of segregated versus aggregated species pairs. The tests did not reveal a large number of significant species pairs in the Vanuatu bird matrix, but in the much smaller Galapagos bird matrix they correctly identified a concentration of segregated species pairs in the genus Geospiza. The Bayesian methods provide for increased selectivity in identifying non-random species pairs, but the analyses will be most powerful if investigators can use a priori biological criteria to identify potential sets of interacting species.

Effects of sap-feeding insect herbivores on growth and reproduction of woody plants: a meta-analysis of experimental studies

Abstract: The majority of generalisations concerning plant responses to herbivory are based on studies of natural or simulated defoliation. However, effects caused by insects feeding on plant sap are likely to differ from the effects of folivory. We assessed the general patterns and sources of variation in the effects of sap feeding on growth, photosynthesis, and reproduction of woody plants through a meta-analysis of 272 effect sizes calculated from 52 papers. Sap-feeders significantly reduced growth (−29%), reproduction (−17%), and photosynthesis (−27%); seedlings suffered more than saplings and mature trees. Deciduous and evergreen woody plants did not differ in their abilities to tolerate damage imposed by sap-feeders. Different plant parts, in particular below- and above-ground organs, responded similarly to damage, indicating that sap-feeders did not change the resource allocation in plants. The strongest effects were caused by mesophyll and phloem feeders, and the weakest by xylem feeders. Generalist sap-feeders reduced plant performance to a greater extent than did specialists. Methodology substantially influenced the outcomes of the primary studies; experiments conducted in greenhouses yielded stronger negative effects than field experiments; shorter (<12 months) experiments showed bigger growth reduction in response to sap feeding than longer experiments; natural levels of herbivory caused weaker effects than infestation of experimental plants by sap-feeders. Studies conducted at higher temperatures yielded stronger detrimental effects of sap-feeders on their hosts. We conclude that sap-feeders impose a more severe overall negative impact on plant performance than do defoliators, mostly due to the lower abilities of woody plants to compensate for sap-feeders’ damage in terms of both growth and photosynthesis.

Estimating parameters of a forest ecosystem C model with measurements of stocks and fluxes as joint constraints

Abstract: We conducted an inverse modeling analysis, using a variety of data streams (tower-based eddy covariance measurements of net ecosystem exchange, NEE, of CO2, chamber-based measurements of soil respiration, and ancillary ecological measurements of leaf area index, litterfall, and woody biomass increment) to estimate parameters and initial carbon (C) stocks of a simple forest C-cycle model, DALEC, using Monte Carlo procedures. Our study site is the spruce-dominated Howland Forest AmeriFlux site, in central Maine, USA. Our analysis focuses on: (1) full characterization of data uncertainties, and treatment of these uncertainties in the parameter estimation; (2) evaluation of how combinations of different data streams influence posterior parameter distributions and model uncertainties; and (3) comparison of model performance (in terms of both predicted fluxes and pool dynamics) during a 4-year calibration period (1997–2000) and a 4-year validation period (“forward run”, 2001–2004). We find that woody biomass increment, and, to a lesser degree, soil respiration, measurements contribute to marked reductions in uncertainties in parameter estimates and model predictions as these provide orthogonal constraints to the tower NEE measurements. However, none of the data are effective at constraining fine root or soil C pool dynamics, suggesting that these should be targets for future measurement efforts. A key finding is that adding additional constraints not only reduces uncertainties (i.e., narrower confidence intervals) on model predictions, but at the same time also results in improved model predictions by greatly reducing bias associated with predictions during the forward run.

Importance of resin ducts in reducing ponderosa pine mortality from bark beetle attack

Abstract: The relative importance of growth and defense to tree mortality during drought and bark beetle attacks is poorly understood. We addressed this issue by comparing growth and defense characteristics between 25 pairs of ponderosa pine (Pinus ponderosa) trees that survived and trees that died from drought-associated bark beetle attacks in forests of northern Arizona, USA. The three major findings of our research were: (1) xylem resin ducts in live trees were >10% larger (diameter), >25% denser (no. of resin ducts mm−2), and composed >50% more area per unit ring growth than dead trees; (2) measures of defense, such as resin duct production (no. of resin ducts year−1) and the proportion of xylem ring area to resin ducts, not growth, were the best model parameters of ponderosa pine mortality; and (3) most correlations between annual variation in growth and resin duct characteristics were positive suggesting that conditions conducive to growth also increase resin duct production. Our results suggest that trees that survive drought and subsequent bark beetle attacks invest more carbon in resin defense than trees that die, and that carbon allocation to resin ducts is a more important determinant of tree mortality than allocation to radial growth.

The impact of protein quality on stable nitrogen isotope ratio discrimination and assimilated diet estimation.

Abstract: Accurately predicting isotopic discrimination is central to estimating assimilated diets of wild animals when using stable isotopes. Current mixing models assume that the stable N isotope ratio (δ15N) discrimination (∆15N) for each food in a mixed diet is constant and independent of other foods being consumed. Thus, the discrimination value for the mixed diet is the combined, weighted average for each food when consumed as the sole diet. However, if protein quality is a major determinant of ∆15N, discrimination values for mixed diets may be higher or lower than the weighted average and will reflect the protein quality of the entire diet and not that of the individual foods. This potential difference occurs because the protein quality of a mixed diet depends on whether, and to what extent, the profiles and amounts of essential amino acids in the individual foods are complementary or non-complementary to each other in meeting the animal’s requirement. We tested these ideas by determining the ∆15N of several common foods (corn, wheat, alfalfa, soybean, and fish meal) with known amino acid profiles when fed singly and in combination to laboratory rats. Discrimination values for the mixed diets often differed from the weighted averages for the individual foods and depended on the degree of complementation. ∆15N for mixed diets ranged from 1.1‰ lower than the weighted average for foods with complementary amino acid profiles to 0.4‰ higher for foods with non-complementary amino acid profiles. These differences led to underestimates as high as 44% and overestimates as high as 36% of the relative proportions of fish meal and soybean meal N, respectively, in the assimilated mixed diets. We conclude that using isotopes to estimate assimilated diets is more complex than often appreciated and will require developing more biologically based, time-sensitive models.

Habitat selection, facilitation, and biotic settlement cues affect distribution and performance of coral recruits in French Polynesia

Abstract: Habitat selection can determine the distribution and performance of individuals if the precision with which sites are chosen corresponds with exposure to risks or resources. Contrastingly, facilitation can allow persistence of individuals arriving by chance and potentially maladapted to local abiotic conditions. For marine organisms, selection of a permanent attachment site at the end of their larval stage or the presence of a facilitator can be a critical determinant of recruitment success. In coral reef ecosystems, it is well known that settling planula larvae of reef-building corals use coarse environmental cues (i.e., light) for habitat selection. Although laboratory studies suggest that larvae can also use precise biotic cues produced by crustose coralline algae (CCA) to select attachment sites, the ecological consequences of biotic cues for corals are poorly understood in situ. In a field experiment exploring the relative importance of biotic cues and variability in habitat quality to recruitment of hard corals, pocilloporid and acroporid corals recruited more frequently to one species of CCA, Titanoderma prototypum, and significantly less so to other species of CCA; these results are consistent with laboratory assays from other studies. The provision of the biotic cue accurately predicted coral recruitment rates across habitats of varying quality. At the scale of CCA, corals attached to the “preferred” CCA experienced increased survivorship while recruits attached elsewhere had lower colony growth and survivorship. For reef-building corals, the behavioral selection of habitat using chemical cues both reduces the risk of incidental mortality and indicates the presence of a facilitator.

Genotypic and environmental variation in specific leaf area in a widespread Alpine plant after transplantation to different altitudes.

Abstract: Specific leaf area (SLA) is an important plant functional trait as it is an indicator of ecophysiological characteristics like relative growth rate, stress tolerance and leaf longevity. Substantial intraspecific variation in SLA is common and usually correlates with environmental conditions. For instance, SLA decreases with increasing altitude, which is understood as adjustment to temperature. It is generally assumed that intraspecific variation is mostly the result of environmentally induced phenotypic plasticity, but genetic effects may also be present, due to local adaptation or genetic drift. In this study, genotypic and environmental effects on SLA were experimentally separated for the widespread Alpine bell flower Campanula thyrsoides by transplanting plants to three common gardens at contrasting altitudes (600, 1,235 and 1,850 m a.s.l.). Seeds were sampled from 18 populations in four phylogeographic regions within the European Alps. A strong plastic response was observed: SLA decreased with increasing altitude of the common gardens (22.0% of variation). The phylogeographic regions were differentiated in SLA in the common gardens (10.1% of variation), indicating that SLA is at least partly genetically determined. Plants from the six easternmost populations experienced a submediterranean climate and showed decreased SLA values in the three common gardens compared to populations to the west, which may be explained as adaptation to drought. Within these submediterranean populations, SLA decreased with altitude of origin in two out of three common gardens. Concluding, SLA shows strong phenotypic plasticity as well as substantial genetic effects, the latter probably being the result of adaptation to local conditions rather than genetic drift.

Immobilizing nitrogen to control plant invasion

Abstract: Increased soil N availability may often facilitate plant invasions. Therefore, lowering N availability might reduce these invasions and favor desired species. Here, we review the potential efficacy of several commonly proposed management approaches for lowering N availability to control invasion, including soil C addition, burning, grazing, topsoil removal, and biomass removal, as well as a less frequently proposed management approach for lowering N availability, establishment of plant species adapted to low N availability. We conclude that many of these approaches may be promising for lowering N availability by stimulating N immobilization, even though most are generally ineffective for removing N from ecosystems (excepting topsoil removal). C addition and topsoil removal are the most reliable approaches for lowering N availability, and often favor desired species over invasive species, but are too expensive or destructive, respectively, for most management applications. Less intensive approaches, such as establishing low-N plant species, burning, grazing and biomass removal, are less expensive than C addition and may lower N availability if they favor plant species that are adapted to low N availability, produce high C:N tissue, and thus stimulate N immobilization. Regardless of the method used, lowering N availability sufficiently to reduce invasion will be difficult, particularly in sites with high atmospheric N deposition or agricultural runoff. Therefore, where feasible, the disturbances that result in high N availability should be limited in order to reduce invasions by nitrophilic weeds.

Locomotor performance in an invasive species: cane toads from the invasion front have greater endurance, but not speed, compared to conspecifics from a long-colonised area

Abstract: Cane toads (Bufo marinus) are now moving about 5 times faster through tropical Australia than they did a half-century ago, during the early phases of toad invasion. Radio-tracking has revealed higher daily rates of displacement by toads at the invasion front compared to those from long-colonised areas: toads from frontal populations follow straighter paths, move more often, and move further per displacement than do toads from older (long-established) populations. Are these higher movement rates of invasion-front toads associated with modified locomotor performance (e.g. speed, endurance)? In an outdoor raceway, toads collected from the invasion front had similar speeds, but threefold greater endurance, compared to conspecifics collected from a long-established population. Thus, increased daily displacement in invasion-front toads does not appear to be driven by changes in locomotor speed. Instead, increased dispersal is associated with higher endurance, suggesting that invasion-front toads tend to spend more time moving than do their less dispersive conspecifics. Whether this increased endurance is a cause or consequence of behavioural shifts associated with rapid dispersal is unclear. Nonetheless, shifts in endurance between frontal and core populations of this invasive species point to the complex panoply of traits affected by selection for increased dispersal ability on expanding population fronts.

Water relations of evergreen and drought-deciduous trees along a seasonally dry tropical forest chronosequence

Abstract: Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (ΨL) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although ΨL did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ18O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar 18O (∆18Ol) and 13C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.

The blind men and the elephant: the impact of context and scale in evaluating conflicts between plant hydraulic safety and efficiency.

Abstract: Given the fundamental importance of xylem safety and efficiency for plant survival and fitness, it is not surprising that these are among the most commonly studied features of hydraulic architecture. However, much remains to be learned about the nature and universality of conflicts between hydraulic safety and efficiency. Although selection for suites of hydraulic traits that confer adequate plant fitness under given conditions is likely to occur at the organismal level, most studies of hydraulic architecture have been confined to scales smaller than the whole plant, such as small-diameter branches and roots. Here we discuss the impact of the spatial and temporal contexts in which hydraulic traits are studied on the interpretation of their role in maintaining plant hydraulic function. We argue that further advances in understanding the ecological implications of different suites of plant hydraulic traits will be enhanced by adopting an integrated approach that considers variation in hydraulic traits throughout the entire plant, dynamic behavior of water transport, xylem tension and water transport efficiency in intact plants, alternate mechanisms that modulate hydraulic safety and efficiency, and alternate measures of hydraulic safety and safety margins.

Non-native grass invasion suppresses forest succession

Abstract: Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.

Biodiversity at the plant-soil interface: microbial abundance and community structure respond to litter mixing.

Abstract: The interactive effects of diversity in plants and microbial communities at the litter interface are not well understood. Mixtures of plant litter from different species often decompose differently than when individual species decompose alone. Previously, we found that litter mixtures of multiple conifers decomposed more rapidly than expected, but litter mixtures that included conifer and aspen litter did not. Understanding the mechanisms underlying these diversity effects may help explain existing anomalous decay dynamics and provide a glimpse into the elusive linkage between plant diversity and the fungi and bacteria that carry out decomposition. We examined the microbial communities on litter from individual plant species decomposing both in mixture and alone. We assessed two main hypotheses to explain how the decomposer community could stimulate mixed-litter decomposition above predicted rates: either by being more abundant, or having a different or more diverse community structure than when microbes decompose a single species of litter. Fungal, bacterial and total phospholipid fatty acid microbial biomass increased by over 40% on both conifer and aspen litter types in mixture, and microbial community composition changed significantly when plant litter types were mixed. Microbial diversity also increased with increasing plant litter diversity. While our data provide support for both the increased abundance hypothesis and the altered microbial community hypothesis, microbial changes do not translate to predictably altered litter decomposition and may only produce synergisms when mixed litters are functionally similar.

Combined effects of precipitation and nitrogen deposition on native and invasive winter annual production in California deserts

Abstract: Primary production in deserts is limited by soil moisture and N availability, and thus is likely to be inXu- enced by both anthropogenic N deposition and precipitation regimes altered as a consequence of climate change. Invasive annual grasses are particularly responsive to increases in N and water availabilities, which may result in competition with native forb communities. Additionally, conditions favoring increased invasive grass production in arid and semi-arid regions can increase Wre risk, negatively impact- ing woody vegetation that is not adapted to Wre. We con- ducted a seeded garden experiment and a 5-year Weld fertilization experiment to investigate how winter annual production is altered by increasing N supply under a range of water availabilities. The greatest production of invasive grasses and native forbs in the garden experiment occurred under the highest soil N (inorganic N after fertilization = 2.99 g m i2 ) and highest watering regime, indicating these species are limited by both water and N. A classiWcation and regression tree (CART) analysis on the multi-year Weld fertilization study showed that winter annual biomass was primarily limited by November-December precipitation. Biomass exceeded the threshold capable of carrying Wre when inorganic soil N availability was at least 3.2 g m i2 in pinon-juniper woodland. Due to water limitation in creo- sote bush scrub, biomass exceeded the Wre threshold only under very wet conditions regardless of soil N status. The CART analyses also revealed that percent cover of invasive grasses and native forbs is primarily dependent on the tim- ing and amount of precipitation and secondarily dependent on soil N and site-speciWc characteristics. In total, our results indicate that areas of high N deposition will be sus- ceptible to grass invasion, particularly in wet years, poten- tially reducing native species cover and increasing the risk of Wre.

A consistent terminology for quantifying species diversity

Abstract: There is a genuine need for consensus on a clear terminology in the study of species diversity given that the nature of the components of diversity is the subject of an ongoing debate and may be the key to understanding changes in ecosystem processes. A recent and thought-provoking paper (Jurasinski et al. Oecologia 159:15–26, 2009) draws attention to the lack of precision with which the terms alpha, beta, and gamma diversity are used and proposes three new terms in their place. While this valuable effort may improve our understanding of the different facets of species diversity, it still leaves us far from achieving a consistent terminology. As such, the conceptual contribution of these authors is limited and does little to elucidate the facets of species diversity. It is, however, a good starting point for an in-depth review of the available concepts and methods.

Differential effects of nutrient-limited primary production on primary, secondary or tertiary consumers

Abstract: Nutritional imbalances between predator and prey are the rule rather than the exception at the lower end of food webs. We investigated the role of different grazers in the propagation of nutritionally imbalanced primary production by using the same primary producers in a three-trophic-level food chain and a four-trophic-level food chain experimental setup. The three-trophic-level food chain consisted of a classic single-cell primary producer (Rhodomonas salina), a metazoan grazer (the copepod Acartia tonsa) and a top predator (the jellyfish Gonionemus vertens), while we added a protozoan grazer (Oxyrrhis marina) as primary consumer to the food chain to establish the four-trophic-level food chain. This setup allowed us to investigate how nutrient-limitation effects change from one trophic level to another, and to investigate the performance of two components of our experimental food chains in different trophic positions. Stoichiometry and fatty acid profiles of the algae showed significant differences between the nutrient-depleted [no N and no P addition (−P), respectively] and the nutrient-replete (f/2) treatments. The differences in stoichiometry could be traced when O. marina was the first consumer. Copepods feeding on these flagellates were not affected by the nutritional imbalance of their prey in their stoichiometry, their respiration rates nor in their developmental rates. In contrast, when copepods were the primary consumer, those reared on the −P algae showed significantly higher respiration rates along with significantly lower developmental rates. In neither of our two experimental food chains did the signals from the base of the food chains travel up to jelly fish, our top predator.

Slow decomposition of lower order roots: a key mechanism of root carbon and nutrient retention in the soil

Abstract: Among tree fine roots, the distal small-diameter lateral branches comprising first- and second-order roots lack secondary (wood) development. Therefore, these roots are expected to decompose more rapidly than higher order woody roots. But this prediction has not been tested and may not be correct. Current evidence suggests that lower order roots may decompose more slowly than higher order roots in tree species associated with ectomycorrhizal (EM) fungi because they are preferentially colonized by fungi and encased by a fungal sheath rich in chitin (a recalcitrant compound). In trees associated with arbuscular mycorrhizal (AM) fungi, lower order roots do not form fungal sheaths, but they may have poorer C quality, e.g. lower concentrations of soluble carbohydrates and higher concentrations of acid-insolubles than higher order roots, thus may decompose more slowly. In addition, litter with high concentrations of acid insolubles decomposes more slowly under higher N concentrations (such as lower order roots). Therefore, we propose that in both AM and EM trees, lower order roots decompose more slowly than higher order roots due to the combination of poor C quality and high N concentrations. To test this hypothesis, we examined decomposition of the first six root orders in Fraxinus mandshurica (an AM species) and Larix gmelinii (an EM species) using litterbag method in northeastern China. We found that lower order roots of both species decomposed more slowly than higher order roots, and this pattern appears to be associated mainly with initial C quality and N concentrations. Because these lower order roots have short life spans and thus dominate root mortality, their slow decomposition implies that a substantial fraction of the stable soil organic matter pool is derived from these lower order roots, at least in the two species we studied.

High mangrove density enhances surface accretion, surface elevation change, and tree survival in coastal areas susceptible to sea-level rise

Abstract: Survival, growth, aboveground biomass accumulation, sediment surface elevation dynamics and nitrogen accumulation in sediments were studied in experimental treatments planted with four different densities (6.96, 3.26, 1.93 and 0.95 seedlings m−2) of the mangrove Rhizophora mucronata in Puttalam Lagoon, Sri Lanka. Measurements were taken over a period of 1,171 days and were compared with those from unplanted controls. Trees at the lowest density showed significantly reduced survival, whilst measures of individual tree growth did not differ among treatments. Rates of surface sediment accretion (means ± SE) were 13.0 (±1.3), 10.5 (±0.9), 8.4 (±0.3), 6.9 (±0.5) and 5.7 (±0.3) mm year−1 at planting densities of 6.96, 3.26, 1.93, 0.95, and 0 (unplanted control) seedlings m−2, respectively, showing highly significant differences among treatments. Mean (±SE) rates of surface elevation change were much lower than rates of accretion at 2.8 (±0.2), 1.6 (±0.1), 1.1 (±0.2), 0.6 (±0.2) and −0.3 (±0.1) mm year−1 for 6.96, 3.26, 1.93, 0.95, and 0 seedlings m−2, respectively. All planted treatments accumulated greater nitrogen concentrations in the sediment compared to the unplanted control. Sediment %N was significantly different among densities which suggests one potential causal mechanism for the facilitatory effects observed: high densities of plants potentially contribute to the accretion of greater amounts of nutrient rich sediment. While this potential process needs further research, this study demonstrated how higher densities of mangroves enhance rates of sediment accretion and surface elevation processes that may be crucial in mangrove ecosystem adaptation to sea-level rise. There was no evidence that increasing plant density evoked a trade-off with growth and survival of the planted trees. Rather, facilitatory effects enhanced survival at high densities, suggesting that managers may be able to take advantage of high plantation densities to help mitigate sea-level rise effects by encouraging positive sediment surface elevation.

How does habitat complexity affect ant foraging success? A test using functional measures on three continents.

Abstract: Habitat complexity can mediate key processes that structure local assemblages through effects on factors such as competition, predation and foraging behaviour. While most studies address assemblage responses to habitat complexity within one locality, a more global approach allows conclusions with greater independence from the phylogenetic constraints of the target assemblages, thus allowing greater generality. We tested the effects of natural and manipulated habitat complexities on ant assemblages from South Africa, Australia and Sweden, in order to determine if there were globally consistent responses in how functional measures of foraging success are regulated by habitat complexity. Specifically, we considered how habitat complexity affected ant foraging rates including the speed of discovery and rate of monopolisation. We also tested if habitat complexity affected the body size index, a size-related morphological trait, of ants discovering resources and occupying and monopolising the resources after 180 min. Ants were significantly slower to discover baits in the more complex treatments, consistent with predictions that they would move more slowly through more complex environments. The monopolisation index was also lower in the more complex treatments, suggesting that resources were more difficult to defend. Our index of ant body size showed trends in the predicted direction for complexity treatments. In addition, ants discovering, occupying and monopolising resources were smaller in simple than in complex natural habitats. Responses of discovering ants to resources in natural habitats were clear in only one of three regions. Consistent with our predictions, habitat complexity thus affected functional measures of the foraging success of ants in terms of measures of discovery and monopolisation rates and body size traits of successful ants. However, patterns were not always equally clear in manipulative and mensurative components of the study.

Natural variation for drought-response traits in the Mimulus guttatus species complex.

Abstract: Soil moisture is a key factor affecting plant abundance and distribution, both across and within species. In response to water limitation, plants have evolved numerous morphological, physiological, and phenological adaptations. In both well-watered and water-limited conditions, we identified considerable natural variation in drought-related whole-plant and leaf-level traits among closely related members of the Mimulus guttatus species complex that occupy a diversity of habitats in the field. The self-fertilizing Mimulus nasutus and serpentine-endemic Mimulus nudatus demonstrated the overall greatest tolerance to soil water limitation, exhibiting the smallest reduction in seed set relative to well-watered conditions. This may be due in part to early flowering, faster fruit development, and low stomatal density. In contrast, flowering of coastal M. guttatus was so delayed that it precluded any seed production in water-limited conditions. This range of phenotypic responses to soil water deficit in Mimulus, coupled with developing genomic resources, holds considerable promise for identifying genomic variation responsible for adaptive responses to soil water availability.

Microbial community shifts influence patterns in tropical forest nitrogen fixation.

Abstract: The role of biodiversity in ecosystem function receives substantial attention, yet despite the diversity and functional relevance of microorganisms, relationships between microbial community structure and ecosystem processes remain largely unknown. We used tropical rain forest fertilization plots to directly compare the relative abundance, composition and diversity of free-living nitrogen (N)-fixer communities to in situ leaf litter N fixation rates. N fixation rates varied greatly within the landscape, and ‘hotspots’ of high N fixation activity were observed in both control and phosphorus (P)-fertilized plots. Compared with zones of average activity, the N fixation ‘hotspots’ in unfertilized plots were characterized by marked differences in N-fixer community composition and had substantially higher overall diversity. P additions increased the efficiency of N-fixer communities, resulting in elevated rates of fixation per nifH gene. Furthermore, P fertilization increased N fixation rates and N-fixer abundance, eliminated a highly novel group of N-fixers, and increased N-fixer diversity. Yet the relationships between diversity and function were not simple, and coupling rate measurements to indicators of community structure revealed a biological dynamism not apparent from process measurements alone. Taken together, these data suggest that the rain forest litter layer maintains high N fixation rates and unique N-fixing organisms and that, as observed in plant community ecology, structural shifts in N-fixing communities may partially explain significant differences in system-scale N fixation rates.