Predicting germination response of four cool-season range grasses to field-variable temperature regimes

Germination of non-dormant seeds under variable-temperature conditions can be predicted from constant-temperature germination data if it is assumed that instantaneous germination rate is independent of thermal history. Thermal-response models of this type have not been validated under simulated field-variable temperature conditions that vary in diurnal pattern, diurnal range and longer-term trends in mean–daily temperature. The purpose of this experiment was to evaluate germination response of thickspike wheatgrass (Elymus lanceolatus), bluebunch wheatgrass (Pseudoroegneria spicata), Sandberg bluegrass (Poa sandbergii) and bottlebrush squirreltail (Elymus elymoides) under both constant and field-variable temperature regimes in the laboratory. It was hypothesized that the thermal history assumption was valid and that constant-temperature data could be used to accurately estimate field-variable temperature response. Seeds were germinated at seven constant temperatures between 5 and 35°C, and under 18 variable-temperature regimes simulating six planting dates at three field sites. Predictions of germination time under variable-temperature conditions were accurate to within a fraction of 1 day up to a cumulative germination percentage of 70% for thickspike wheatgrass, 60% for bluebunch wheatgrass, 55% for Sandberg bluegrass and 70% for bottlebrush squirreltail. It was concluded that, for the variable-temperature regimes tested in this experiment, the thermal-history assumption was valid for earlier-germinating subpopulations.     

Do invasive plants structure microbial communities to accelerate decomposition in intermountain grasslands?

Invasive plants are often associated with greater productivity and soil nutrient availabilities, but whether invasive plants with dissimilar traits change decomposer communities and decomposition rates in consistent ways is little known. We compared decomposition rates and the fungal and bacterial communities associated with the litter of three problematic invaders in intermountain grasslands; cheatgrass (Bromus tectorum), spotted knapweed (Centaurea stoebe) and leafy spurge (Euphorbia esula), as well as the native bluebunch wheatgrass (Pseudoroegneria spicata). Shoot and root litter from each plant was placed in cheatgrass, spotted knapweed, and leafy spurge invasions as well as remnant native communities in a fully reciprocal design for 6 months to see whether decomposer communities were species‐specific, and whether litter decomposed fastest when placed in a community composed of its own species (referred to hereafter as home‐field advantage–HFA). Overall, litter from the two invasive forbs, spotted knapweed and leafy spurge, decomposed faster than the native and invasive grasses, regardless of the plant community of incubation. Thus, we found no evidence of HFA. T‐RFLP profiles indicated that both fungal and bacterial communities differed between roots and shoots and among plant species, and that fungal communities also differed among plant community types. Synthesis. These results show that litter from three common invaders to intermountain grasslands decomposes at different rates and cultures microbial communities that are species‐specific, widespread, and persistent through the dramatic shifts in plant communities associated with invasions.     

Response of early and late semiarid seral species to nitrogen and phosphorus gradients

Above- and below-ground biomass production, nitrogen (N) and phosphorus (P) tissue concentrations, and root: shoot ratios were examined for five species that are characteristic of a semiarid successional sequence under controlled greenhouse conditions. In two simultaneous experiments, seedlings of one forb, two grass, and two shrub species important in a sagebrush successional sere, were subjected to seven levels of N and P. Results of the experiments suggest distinct differences in nutrient response patterns between early and late seral species. Early seral species produced more biomass but had lower tissue nutrient concentrations than late seral species. As N and P availabilities decreased, late seral species displayed characteristics indicative of increasing competitive advantage over those of early seral species. Root: shoot ratios of the five species primarily reflected patterns related to lifeform, but with some early and late seral characteristics. Results from this study 1) confirm that nutrient use pattern, nutrient availability, and seral position relationships characteristic of mesic ecosystems hold equally true for semiarid systems, and 2) suggest that nutrients are important organizing factors in semiarid ecosystems.     

Can R*s Predict Invasion in Semi-arid Grasslands?

We estimated R*s and tested the applicability of R* theory on nonindigenous plant invasions in semi-arid rangeland. R* is the concentration of a resource that a species requires to survive in a habitat. R* theory predicts that a species with a lower R* for the most limiting resource will competitively displace a species with a higher R* under equilibrium conditions. In a greenhouse, annual sunflower (Helianthus annuus L.), bluebunch wheatgrass (Agropyron spicatum Pursh), and spotted knapweed (Centaurea maculosa Lam.) were grown in monoculture and 2- and 3-species mixtures for three growth periods in an attempt to reduce soil NO₃-N concentrations below each species’ R*. At the end of each growth period, aboveground biomass by species and soil plant available nitrogen were sampled. Decreasing biomass coupled with decreasing soil plant available nitrogen was used to quantify R*s for the three species. R*s for annual sunflower, bluebunch wheatgrass, and spotted knapweed were estimated to be 0.6±0.16 ppm NO₃⁻, less than 0.05 ppm NO₃⁻, and 0.6±0.13 ppm NO₃⁻, respectively. Estimated R*s did not predict the outcome of competition among species. To successfully predict plant community dynamics on semi-arid rangeland with and without the presence of a nonindigenous invasive species, a more comprehensive model that includes mechanisms in addition to competition may have to be considered. We speculate that R* theory may prove most useful for predicting the outcome of competition within functional groups.     

Germination and Emergence of Primed Grass Seeds Under Field and Simulated-field Temperature Regimes

Seed priming may enhance establishment success of cool-seasonrange grasses which must compete with annual weeds for earlyspring moisture. Previous priming studies have confirmed germinationrate enhancement for these species but relative treatment effectsunder field-temperature conditions have not been assessed. Weprimed seeds of thickspike wheatgrass [Elymus lanceolatus(Scribn.and J. G. Smith) Gould], bluebunch wheatgrass [Pseudoroegneriaspicata(Pursh) Löve], Sandberg bluegrass (Poa sandbergiiVasey.) and bottlebrush squirreltail [Elymus elymoides(Raf.)Swezey] and evaluated their relative emergence rate in threesoil types as a function of spring-planting date. Germinationresponse was simultaneously evaluated in laboratory germinatorsthat were programmed to simulate the field-temperature regimeat planting depth. Seed priming enhanced both germination andemergence rate with the greatest effect occurring during theearlier, cooler planting dates. Total emergence and emergencerate in the field were lower than for the equivalent germinationresponse in the laboratory. Thermal-germination response wasmodelled and predictions developed for evaluating potentialgermination under late winter/early spring soil-temperatureregimes. Modelling results predicted that greater germinationenhancement would have been possible at earlier planting datesthan were measured in the field experiment.Copyright 2000 Annalsof Botany Company Bunchgrass, germination, emergence, priming, rate, temperature     

Competition along a nutrient gradient: A case study with Daucus carota and Chenopodium album

A binary competition experiment between carrot (Daucus carota L.) and Chenopodium album L. was conducted in a greenhouse at seven population densities and eight nutrient concentrations to investigate the effects of a nutrient gradient on plant competition in both monocultures and mixtures. The patterns of carrot biomass allocation (measured as root : shoot ratio) in monocultures and mixtures with C. album were affected by both nutrient availability and population density. Chenopodium album had a broader response to nutrient concentrations than carrot. The maximum yield of carrot in both monocultures and mixtures occurred at fourfold the standard concentration of nutrients, while C. album in both monocultures and mixtures had the maximum yield at 16-fold the standard nutrient concentration. The yield–density relationship of carrot tended to be increasing or asymptotic at lower nutrient concentrations but parabolic at higher concentrations, whereas that of C. album was little affected by nutrient availability. Nutrient availability had a profound influence on the competitive relationships between the two species: at both low and high nutrient concentrations, C. album tended to be more competitive than carrot, while at intermediate levels of nutrients, carrot was more competitive than C. album. Our results suggest that in relation to competitive performance, the weed has a greater ability to adapt itself to varying environments than does the crop. Additionally, the relative merits of the quantitative measures of competitive ability are briefly discussed.     

Arbuscular mycorrhizal fungi enhance spotted knapweed growth across a riparian chronosequence

Arbuscular mycorrhizal fungi (AMF) mediate nutrient uptake that accelerates plant growth and reproduction. Thus, AMF may promote plant invasions often observed along rivers. We assessed the importance of AMF in improving growth of the invasive species, spotted knapweed (Centaurea stoebe), during succession of riparian vegetation along a flood plain in Montana, USA. We grew spotted knapweed with and without AMF in soils collected from riparian sites ranging from 1 to 72 years old and measured the plant’s growth response to AMF. We observed variability in relative effects of AMF, with greatest growth benefits in recently deposited alluvial sediments. We then separated effects of soil and inoculum source by growing spotted knapweed with soils and inocula collected from young or old sites and found that growth responses were greatest in young soils regardless of inoculum source. Our results demonstrate that AMF directly benefit growth of spotted knapweed, especially in soils that typify early successional sites on this alluvial flood plain.     

Additive effects of aboveground and belowground herbivores on the dominance of spotted knapweed (Centaurea stoebe)

Spotted knapweed (Centaurea stoebe) is found in over 3 million ha of rangeland and forests across North America, and evidence supporting the use of biological control as a regional method to reduce infestations and their associated impacts remains inconclusive. Several species of insects have been reported to reduce plant densities in some areas; however, rigorous studies that test combinations of these species and the influence of resource availability are lacking. We examined the singular and combined effects of herbivory by a root weevil (Cyphocleonus achates) and a flower head weevil (Larinus minutus) on the growth and flower production of C. stoebe. We also manipulated soil resource fertility as an additional factor that could explain the outcomes of contradictory biological control herbivore effects on C. stoebe. In a greenhouse study, herbivory by C. achates decreased flower production for plants across all resource environments. In a caged common garden study, the negative effects of herbivory also did not interact with soil nutrient status. However, the presence of plant competition further decreased knapweed growth, and the negative effects of concurrent herbivory by C. achates and L. minutus on plant biomass and flower production were additive. Derived within the context of variable levels of soil nutrient availability and competing vegetation, these results support the cumulative stress hypothesis and the contention that combined above- and belowground herbivory can reduce spotted knapweed densities and reduce the ecological and economic impacts of this species in rangelands of western North America.     

Morphological plasticity following species-specific recognition and competition in two perennial grasses

Morphological characteristics and biomass allocation of two perennial grasses, Pseudoroegneria spicata (Pursh) A. Löve ssp. spicata (bluebunch wheatgrass) and Agropyron desertorum (Fisch. ex Link) Schult. (crested wheatgrass), were compared under different competition and nutrient treatments. The competitive responses of two plants grown in containers under field conditions were assessed in monocultures and mixtures in two experiments using different scales of nutrient application. In the Small-Scale Experiment, a localized fertilization was applied in the rooting zone between two plants; in the Large-Scale Experiment the entire container was supplied with nutrients. Agropyron responded more vigorously to fertilization than did Pseudoroegneria, but based on the relative performance of Agropyron in monoculture and mixture, it was not superior to Pseudoroegneria in resource competition. Pseudoroegneria was apparently able to recognize neighboring plants as either conspecifics or individuals of the other species. The responses included changes in shoot architecture, root morphology, and allocation between roots and shoots. Agropyron generally did not exhibit such morphological flexibility. In field plot plantings of 4-yr-old tussocks similar shoot differences were seen in Pseudoroegneria. There was, however, no indication of superior resource competition for Agropyron. Thus, any early advantage of Agropyron in vigorous growth of young plants in response to nutrients was apparently lost by the time the plants had reached this stage of development. Morphological and allocation flexibility of Pseudoroegneria may have compensated for slower, less vigorous growth. If species-specific recognition and morphological plasticity are common in nature, this complicates our attempts to understand mechanisms of competition.     

A knapweed biological control perspective

Diffuse and spotted knapweed (Centaurea diffusa Lam. and C. stoebe micranthos (Gugler) Hayek) are Eurasian plants that devastate dry and mesic North American grasslands. They have a mutualistic association with arbuscular mycorrhizal fungal (AMF) phylotypes with hyphal links to nearby plants and a nutrient flux to the strongest sink, usually knapweed. They displace many AMF beneficial to grass and affect knapweed nutrient allocation, biology, knapweed insects and probably root necrosis and emergence of ant buried seed. AMF determined nutrient root or shoot allocation determines nutrient shoot and root allocation and the benefit to root or seed-head insect species and whether C. diffusa is an annual–biannual or a semelparous perennial needing 5 or more years to flower. Both knapweeds do well without its AMF phylotypes without competition in fertile soil. In grass in Eurasia, they have a community of seven seed-head species segregated by head development stage. Prolonged seed dormancy buffered knapweed decline that resulted in release of a surfeit seed-head species. The presence of an eliasome on the seed and vigorous seedling clumps suggests burial by myrmecochorous ants with AMF supplied carbon supporting their growth. The root species community is segregated by habitat, climate, root part, and size. With larval induced compensatory growth and AMF nutrient sharing, the growth of plants with and without a larva was the same. On feeding completion, a nutrient out flux from the attacked plants reduced growth; but without killing. This needs a dual species or a repeated single species attack. Root species packing increases knapweed utilization; but the four approved species are insufficient for maximum utilization. Two additions are suggested. The aim of the paper is to provide enough understanding of the AMF and its plant and insect interactions to facilitate knapweed biological control and avoid past mistakes.     

Is spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis> L.) patch size related to the effect on soil and vegetation properties?

Spotted knapweed (Centaurea stoebe L. subsp. Micranthos (Gugler) Hayek) was first introduced in the 1890s from Europe into western North America, where it now occupies over three million hectares of rangeland and pasture in 14 states and two Canadian provinces, reducing forage production and causing economic damage. Despite many reported effects spotted knapweed can have on soils and native vegetation, it is not known whether patch size is correlated with these ecosystem-level effects. The objective of our study was to determine whether the effects of spotted knapweed on plant composition and soil properties was related to spotted knapweed patch size. We asked the following questions: (1) Are there differences in plant species richness and diversity between small and large knapweed patches? and (2) Do soil water and soil mineral nutrient properties change depending on knapweed patch size? Twenty-four knapweed patches, and paired natural grassland plots, were randomly selected within Lac du Bois Provincial Park, British Columbia, Canada. Knapweed patch size ranged from 6 to 366 m². Sampling and analysis revealed a significant effect of knapweed patch size on soil and vegetation properties. Soil P, soil temperature, and total dry plant biomass (g/0.25 m²) increased, while soil N, soil C, and soil moisture decreased with patch size. Since our results show that spotted knapweed patch size is related to degree of soil alteration, it is important to consider size of patch when modeling the impact of spotted knapweed in North America. Since large patches of spotted knapweed seem to have a proportionately greater effect on soil chemistry properties, large patches may move the system further away from a point where it is possible to restore the site to pre-invasion conditions.     

Effects of plant competition,seed predation,and nutrient limitation on seedling survivorship of spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis>)

We measured seed germination and seedling survivorship of spotted knapweed, Centaurea stoebe, in a series of laboratory and field experiments to evaluate the efficacy of seed limitation as a management focus. This work was initiated 6 years after introduction of several biological control agents. The soil seed bank of the site used in this study contained a mean density of 5,848 seeds/m² (ranging from 0 to 16,364 seeds/m²), and 92% of the seeds isolated from soils were shriveled, discolored, and/or partially decayed. Additionally, none of the intact seeds germinated, suggesting that the viable seed bank at our field study site has been exhausted. Centaurea stoebe seeds were planted into pots under a range of soil nitrogen (N) availability, with half of the pots containing a single density of previously established seedlings of a native cool-season grass, slender wheatgrass (Elymus trachycaulus). A watering regime mimicking local precipitation was applied. Spotted knapweed exhibited large biomass responses to N addition, but the presence of grasses suppressed the ability to exploit this N. Surprisingly, low soil N conditions improved knapweed survivorship in the presence of grasses. Nevertheless, recruitment and biomass were still far below the levels reached in the absence of competition. To evaluate the effect of density on successful recruitment, Centaurea stoebe seed was introduced into a meadow at three densities matching reduced levels of seed production under the constraints of seed predators. These densities were sown with or without a seed mixture of native species, into an existing plant community lacking C. stoebe, and seedling recruitment was recorded over 2.5 years. Across all plots and densities sown (568–2,272 seeds m⁻² year⁻¹), seedling recruitment was less than 1%. The invasion potential of spotted knapweed was greatly diminished when realistic levels of plant competition and biological control limit seed production. We therefore conclude that a combination of seed limitation and shortage of ‘safe sites’ within undisturbed vegetation can limit densities of C. stoebe.     

Root traits associated with nutrient exploitation following defoliation in three coexisting perennial grasses in a semi-arid savanna

Experiments were conducted to evaluate root traits associated with nutrient exploitation following defoliation in three coexisting perennial grasses in a semi‐arid savanna. Root length density was determined within soil cores directly beneath plants, nitrogen uptake was evaluated by excised‐root assay with (¹⁵NH₄)₂SO₄, and mycorrhizal root colonization was estimated by observation of root segments. Root length density was lowest for Bouteloua curtipendula, intermediate for Eriochloa sericea, and highest for Aristida purpurea indicating that root length density was a more important trait for the mid‐seral than the late‐seral species. Rates of ¹⁵N uptake were greatest in the least grazing tolerant late‐seral species, E. sericea, intermediate in the mid‐seral species, A. purpurea, and lowest in the most grazing tolerant late‐seral species, B. curtipendula. Two successive defoliations reduced ¹⁵N uptake 60% in the late‐seral species with the greatest uptake rate (E. sericea), but not in species with lowest uptake rates (B. curtipendula). Root length colonization was consistently high (33–61%) in all three species suggesting that these C₄ perennial grasses may function as obligate mycotrophs. Contrasting responses among the two late‐seral species indicate that the least grazing tolerant species, E. sericea, appears best adapted for nutrient exploitation while the most grazing tolerant species, B. curtipendula, appears best adapted for efficient nutrient retention. Contrasting responses of nitrogen uptake to short‐term defoliation parallel the population responses of these two coexisting late‐seral species to long‐term herbivory. These data indicate that herbivory may shift interspecific competitive interactions by mediating nutrient exploitation and that a trade‐off may exist between nutrient exploitation and herbivory tolerance in these species.     

Allometry of root branching and its relationship to root morphological and functional traits in three range grasses

The study of proportional relationships between size, shape, and function of part of or the whole organism is traditionally known as allometry. Examination of correlative changes in the size of interbranch distances (IBDs) at different root orders may help to identify root branching rules. Root morphological and functional characteristics in three range grasses {bluebunch wheatgrass [Pseudoroegneria spicata (Pursh) L?ve], crested wheatgrass [Agropyron desertorum (Fisch. ex Link) Schult.×A. cristatum (L.) Gaert.], and cheatgrass (Bromus tectorum L.)} were examined in response to a soil nutrient gradient. Interbranch distances along the main root axis and the first-order laterals as well as other morphological and allocation root traits were determined. A model of nutrient diffusivity parameterized with root length and root diameter for the three grasses was used to estimate root functional properties (exploitation efficiency and exploitation potential). The results showed a significant negative allometric relationship between the main root axis and first-order lateral IBD (P ≤ 0.05), but only for bluebunch wheatgrass. The main root axis IBD was positively related to the number and length of roots, estimated exploitation efficiency of second-order roots, and specific root length, and was negatively related to estimated exploitation potential of first-order roots. Conversely, crested wheatgrass and cheatgrass, which rely mainly on root proliferation responses, exhibited fewer allometric relationships. Thus, the results suggested that species such as bluebunch wheatgrass, which display slow root growth and architectural root plasticity rather than opportunistic root proliferation and rapid growth, exhibit correlative allometry between the main axis IBD and morphological, allocation, and functional traits of roots.     

Nutrient utilization in Quercus leucotrichophora and Pinus roxburghii seedlings at five soil fertility levels

Abstract. Nutrient utilization and growth performance of Pinus roxburghii Sarg. and Quercus leucotrichophora A. Camus, the two major forest trees of the Central Himalaya Mountains between 1000 and 2000 m elevation, were analysed at different nutrient levels. The early successional P. roxburghii occurs naturally on nutrient-poor sites and the late successional Q. leucotrichophora on nutrient-rich sites. Seedlings of these two species were grown both in monoculture and mixed, in plastic bags containing 1 kg soil and representing five levels of nutrients (NPK) in a 12 : 32 : 16 ratio. The P. roxburghii seedlings showed significantly greater nutrient extraction efficiency (nutrient unit extracted by seedlings per nutrient unit in the soil) than the Q. leucotrichophora seedlings, and this difference was increased in interspecific competition, particularly at the higher nutrient levels. In interspecific competition the extraction efficiencies of P. roxburghii for N, P and K were distinctly higher than those of Q. leucotrichophora. This we consider to be the main reason for the replacement of Q. leucotrichophora by P. roxburghii, even on nutrient-rich soils, after disturbance and subsequent increase in light availability. In both the species, re-translocation of nutrients from senescing leaves declined with increasing soil fertility. For example, re-translocation for P. roxburghii was 49% N, 30% P and 32% K at the lowest soil fertility, and 20% N, 8% P and 13% K at the highest soil fertility.     

A comparison of phenotypic plasticity between two species occupying different positions in a successional sequence

We compared the phenotypic plasticity of two greenhouse-grown species (Corispermum macrocarpum and Salsola collina) occupying different positions in a successional sequence in Horqin Sandy Land, by treating with different population density and the availability of soil nutrients and water. The same species can exhibit different patterns of plasticity in response to different environmental factors. In the soil nutrient treatments, the plasticity pattern of S. collina could be described as “master-of-some”. However, in the soil-water and population-density treatments, it showed no significant difference from C. macrocarpum in the reaction norm for plasticity. It was similar to a “jack-of-all-trades” plasticity pattern. Contrary to the previous conclusion that late successional species had higher reproductive allocation than early successional species, in this successional sequence, the late species had lower reproductive allocation in all treatments. Reproductive allocation of both species increased with the increase in water availability and also increased with a decrease in nutrient levels. However, density had no effect on reproductive allocation. Although the root:shoot ratio increased with decreasing water availability, there were no differences in the plasticity pattern for this trait in both species. Root:shoot ratio was, however, not significantly affected by nutrient availability and density. In a word, the plasticity patterns of invaders are adapted to the analyses of succession.     

Effects of a nutrient pulse supply on nutrient status of the Mediterranean trees Quercus ilex subsp. ballota and Pinus halepensis on different soils and under different competitive pressure

Nutrient availability is a key factor in Mediterranean ecosystems that affects the primary productivity and the community structure. The great variability of its natural availability is now increasing due to frequent fires, pollution events and changes in rainfall regime associated to climate change. Quercus ilex ssp. ballota and Pinus halepensis are the most abundant tree species in the NW Mediterranean basin. They frequently compete in the early and middle successional stages. We investigated the effects of N and P pulse supplies on nutrient uptake capacity in these two species in an after-fire field area and in nursery conditions on different soil types and competing conditions. In the field, N fertilisation had weak effects on nutrient concentration and mineralomass likely as a consequence of this nutrient not being limiting in this field site whereas P fertilisation increased the P mineralomass and the Mg, S, Fe, K and Ca concentrations and mineralomass in the different biomass fractions of both species 1 and 3 years after fertilisation application. In the nursery experiments, P fertilisation increased the mineralomass and concentrations of P, Mg, S, Fe, K and Ca in all biomass fractions including the roots in both species and in different soils and competition conditions. The increment of nutrient mineralomass was due to both the increase of growth and of nutrient concentrations. Both species were able to absorb significant amounts of the P applied by fertilisation (between 5 and 20%) in short time (18 months). Competing vegetation decreased the positive effects of P fertilisation, and in many cases the negative effect of competing vegetation on nutrient mineralomass was stronger when P availability was increased by fertilisation. Q. ilex subsp ballota showed a greater competitive ability for P than the more pioneer species Pinus halepensis in the field but not in the nursery conditions. Pinus halepensis had greater nutrient mineralomass in calcareous than in siliceous soils. Q. ilex subsp. ballota had a higher root biomass allocation and root nutrient allocation than P. halepensis, but both species showed a high capacity to increase their nutrient uptake when its availability increased by fertilisation, thus assuring a great nutrient reserve for future growth periods and contributing to retain nutrients in the ecosystem.     

Nutrient conservation strategies in native Andean‐Patagonian forests

Abstract. Nutrient conservation in vegetation affects rates of litter decomposition and soil nutrient availability. Although resorption has been traditionally considered one of the most important plant strategies to conserve nutrients in temperate forests, long leaf life‐span and low nutrient requirements have been postulated as better indicators. We aimed at identifying nutrient conservation strategies within characteristic functional groups of NW Patagonian forests on Andisols. We analysed C‐, N‐, P‐, K‐ and lignin‐concentrations in mature and senescent leaves of ten native woody species within the functional groups: broad‐leaved deciduous species, broad‐leaved evergreens and conifers. We also examined mycorrhizal associations in all species. Nutrient concentration in mature leaves and N‐ resorption were higher in broad‐leaved deciduous species than in the other two functional groups. Conifers had low mature leaf nutrient concentrations, low N‐resorption and high lignin/N ratios in senescent leaves. P‐ and K‐resorptions did not differ among functional groups. Broad‐leaved evergreens exhibited a species‐dependent response. Nitrogen in mature leaves was positively correlated with both N resorption and soil N‐fertility. Despite the high P‐retention capacity of Andisols, N appeared to be the more limiting nutrient, with most species being proficient in resorbing N but not P. The presence of endomycorrhizae in all conifers and the broad‐leaved evergreen Maytenus boaria, ectomycorrhizae in all Nothofagus species (four deciduous, one evergreen), and cluster roots in the broad‐leaved evergreen Lomatia hirsuta, would be possibly explaining why P is less limiting than N in these forests.     

Nutrients in fruits as determinants of resource tracking by birds

Fruit pulp is an important source of nutrients for many bird species. Fruit‐eating birds use a variety of strategies to cope with changes in the availability of fruits, exhibiting a remarkable ability to track resources. We assessed the role of nutrient availability in the fruiting environment as a factor driving resource tracking by fruit‐eating birds. Fruit consumption by the four most common frugivorous species in a 6‐ha plot in the Southern Yungas montane forest of Argentina was assessed. We determined the content of selected nutrients (soluble carbohydrates, proteins, phenols, ascorbic acid and essential minerals) in 22 fruiting plant species eaten by birds, and measured fruit–frugivore interactions and the availability of nutrients and dry fruit pulp mass over 2 years. There was strong temporal covariation in the availability of the selected nutrients in fruits across the study period. Similarly, the availability of nutrients in the fruiting environment covaried with pulp mass. Fruit consumption by the four commonest bird species and the abundance of most species were positively associated with nutrient availability and dry pulp mass. Nutrient availability was a good predictor of temporal fruit tracking by three of the four commonest frugivores. Despite large differences in particular nutrient concentrations in fruits, overall nutrient (and pulp) quantity in the fruiting environment played a greater role in fruit tracking than did the nutritional quality of individual fruits. While overall nutrient availability (i.e. across fruit) and total pulp mass were important determinants of fruit tracking, we suggest that plant species‐specific differences in fruit nutrient concentration may be important in short‐term foraging decisions involved in fruit choice and nutritional balance of birds.     

Invasion of an exotic forb impacts reproductive success and site fidelity of a migratory songbird

Although exotic plant invasions threaten natural systems worldwide, we know little about the specific ecological impacts of invaders, including the magnitude of effects and underlying mechanisms. Exotic plants are likely to impact higher trophic levels when they overrun native plant communities, affecting habitat quality for breeding songbirds by altering food availability and/or nest predation levels. We studied chipping sparrows (Spizella passerina) breeding in savannas that were either dominated by native vegetation or invaded by spotted knapweed (Centaurea maculosa), an exotic forb that substantially reduces diversity and abundance of native herbaceous plant species. Chipping sparrows primarily nest in trees but forage on the ground, consuming seeds and arthropods. We found that predation rates did not differ between nests at knapweed and native sites. However, initiation of first nests was delayed at knapweed versus native sites, an effect frequently associated with low food availability. Our seasonal fecundity model indicated that breeding delays could translate to diminished fecundity, including dramatic declines in the incidence of double brooding. Site fidelity of breeding adults was also substantially reduced in knapweed compared to native habitats, as measured by return rates and shifts in territory locations between years. Declines in reproductive success and site fidelity were greater for yearling versus older birds, and knapweed invasion appeared to exacerbate differences between age classes. In addition, grasshoppers, which represent an important prey resource, were substantially reduced in knapweed versus native habitats. Our results strongly suggest that knapweed invasion can impact chipping sparrow populations by reducing food availability. Food chain effects may be an important mechanism by which strong plant invaders impact songbirds and other consumers.