Showing papers in "Oecologia in 1995"

The promotive effect of smoke derived from burnt native vegetation on seed germination of Western Australian plants

Abstract: Exposure of dormant seed to cold smoke derived from burnt native vegetation had a positive influence on germination in one or more seed provenances in 45 out of 94 species of native Western Australian plants that are normally hard to germinate. When tested under controlled conditions some species showed earlier germination in smoke treatments than controls; in others smoke-treated seeds continued to germinate for several weeks after controls had achieved full germination. In the remainder, treated and control seeds germinated to similar time schedules. A group of 23 species which responded positively had previously been recorded as extremely difficult or impossible to germinate using conventional techniques. These included members of the genera Geleznowia (Rutaceae), Hibbertia (Dilleniaceae), Stirlingia (Proteaceae), Verticordia (Myrtaceae), Actinostrobus (Cupressaceae) and Pimelea (Thymelaeaceae). Both large- and small-seeded species were encountered amongst the positively responding taxa, which encompassed representatives of 15 families and 26 genera of dicotyledons, 5 families and 8 genera of monocotyledons and the gymnosperm Actinostrobus acuminatus. Sowing seeds on smoke-fumigated filter papers or watering with aqueous eluates of smoke elicited similar degrees of stimulation of germination, as did exposure to gaseous smoke in a readily germinating species Anigozanthos manglesii (Haemodoraceae) and the normally intractable species Lysinema ciliatum (Epacridaceae). Exposing recently burnt and unburnt natural bushland sites to smoke, smoked water or smoked dry sand elicited a significant germination response in 15 species. Over one third of the species sampled in the burnt site exhibited germination additional to that caused by the fire. Data are discussed in relation to previous germination studies on Australian and other taxa.

Different photosynthesis-nitrogen relations in deciduous hardwood and evergreen coniferous tree species.

Abstract: The relationship between photosynthetic capacity (A max) and leaf nitrogen concentration (N) among all C3 species can be described roughly with one general equation, yet within that overall pattern species groups or individual species may have markedly different A max-N relationships. To determine whether one or several predictive, fundamental A max-N relationships exist for temperate trees we measured A max, specific leaf area (SLA) and N in 22 broad-leaved deciduous and 9 needle-leaved evergreen tree species in Wisconsin, United States. For broad-leaved deciduous trees, mass-based A max was highly correlated with leaf N (r 2=0.75, P 0.25) in the evergreen conifers. Compared to deciduous trees at a common leaf N (mass or area basis), evergreen trees had lower A max and SLA. For all data pooled, both leaf N and A max on a mass basis were correlated (r 2=0.6) with SLA; in contrast, area-based leaf N scaled tightly with SLA (r 2=0.81), but area-based A max did not (r 2=0.06) because of low A max per unit N in the evergreen conifers. Multiple regression analysis of all data pooled showed that both N (mass or area basis) and SLA were significantly (P<0.001) related to A max on mass (r 2=0.80) and area (r 2=0.55) bases, respectively. These results provide further evidence that A max-N relationships are fundamentally different for ecologically distinct species groups with differing suites of foliage characteristics: species with long leaf life-spans and low SLA, whether broad-leaved or needle-leaved, tend to have lower A max per unit leaf N and a lower slope and higher intercept of the A max-N relation than do species with shorter leaf life-span and higher SLA. A single global A max-N equation overestimates and underestimates A max for temperate trees at the upper and lower end of their leaf N range, respectively. Users of A max-N relationships in modeling photosynthesis in different ecosystems should appreciate the strengths and limitations of regression equations based on different species groupings.

Determination of the quantum efficiency of photosystem II and of non-photochemical quenching of chlorophyll fluorescence in the field

Abstract: A newly developed portable chlorophyll fluorometer in combination with a special leaf clip holder was used for assessing photosynthetic activity of attached sun leaves of Fagus sylvatica and Cucurbita pepo under field conditions. During diurnal time courses, fluorescence yield, photosynthetic photon flux density (PPFD) incident on the leaf plane, and leaf temperature were measured and quantum efficiency of photosystem II (PS II), apparent relative electron transport rates, and non-photochemical fluorescence quenching (NPQ) calculated. In both species, quantum efficiency followed closely the incident PPFD and no hysteresis could be observed during the day. Apparent electron transport rate showed light saturation above a PPFD of 700 μmol m−2 s−1 in F. sylvatica, while in C. pepo no saturation was visible up to 1400 μmol m−2 s−1. NPQ was closely correlated to excessive PPFD calculated from the PS II quantum yield. Maximal NPQ observed was 3.3 Although the beech leaf was exposed for a considerable time to PPFD values of 1400–1500 μmol m−2 s−1 and leaf temperatures between 30 and 35°C, no obvious signs for sustained photodamage could be observed. The data demonstrate the potential of chlorophyll fluorescence measurements to analyse photosynthetic performance under field conditions with minimal disturbance of the plant. Potential error sources due to the geometry of the leaf clip holder used are discussed.

External hyphal production of vesicular-arbuscular mycorrhizal fungi in pasture and tallgrass prairie communities.

Abstract: External hyphae of vesicular-arbuscular mycorrhizal (VAM) fungi were quantified over a growing season in a reconstructed tallgrass prairie and an ungrazed cool-season pasture. In both sites, hyphal lengths increased throughout the growing season. Peak external hyphal lengths were 111 m cm−3 of soil in the prairie and 81 m cm−3 of soil in the pasture. These hyphal lengths calculate to external hyphal dry weights of 457 μg cm−3 and 339 μg cm−3 of soil for prairie and pasture communities, respectively. The relationships among external hyphal length, root characteristics, soil P and soil moisture were also determined. Measures of gross root morphology [e.g., specific root length (SRL) and root mass] have a strong association with external hyphal length. Over the course of the study, both grassland communities experienced a major drought event in late spring. During this period a reduction in SRL occurred in both the pasture and prairie without a measured reduction in external hyphal length. Recovery for both the pasture and prairie occurred not by increasing SRL, but rather by increasing external hyphal length. This study suggests that growth is coordinated between VAM hyphae and root morphology, which in turn, are constrained by plant community composition and soil nutrient and moisture conditions.

Fire in the Brazilian Amazon: 1. Biomass, nutrient pools, and losses in slashed primary forests.

Abstract: Deforestation in the Brazilian Amazon has resulted in the conversion of >230,000 km2 of tropical forest, yet little is known on the quantities of biomass consumed or the losses of nutrients from the ecosystem. We quantified the above-ground biomass, nutrient pools and the effects of biomass burning in four slashed primary tropical moist forests in the Brazilian Amazon. Total above-ground biomass (TAGB) ranged from 292 Mg ha-1 to 436 Mg ha-1. Coarse wood debris (>20.5 cm diameter) was the dominant fuel component. However, structure of the four sites were variable. Coarse wood debris comprised from 44% to 69% of the TAGB, while the forest floor (litter and rootmat) comprised from 3.7 to 8.0% of the TAGB. Total biomass consumption ranged from 42% to 57%. Fires resulted in the consumption of >99% of the litter and rootmat, yet <50% of the coarse wood debirs. Dramatic losses in C, N, and S were quantified. Lesser quantities of P, K, and Ca were lost by combustion processes. Carbon losses from the ecosystem were 58–112 Mg ha-1. Nitrogen losses ranged from 817 to 1605 kg ha-1 and S losses ranged from 92 to 122 kg ha-1. This represents losses that are as high as 56%, 68%, and 49% of the total above-ground pools of these nutrients, respectively. Losses of P were as high as 20 kg ha-1 or 32% of the above-ground pool. Losses to the atmosphere arising from primary slash fires were variable among sites due to site differences in concentration, fuel biomass, and fuel structure, climatic fluctuations, and anthropogenic influences. Compared to fires in other forest ecosystems, fires in slashed primary tropical evergreen forests result in among the highest total losses of nutrients ever measured. In addition, the proportion of the total nutrient pool lost from slash fires is higher in this ecosystem compared to other ecosystems due to a higher percentage of nutrients stored in above-ground biomass.

Interaction of pre-attack and induced monoterpene concentrations in host conifer defense against bark beetle-fungal complexes.

Abstract: Two pine species (Pinus resinosa, P. banksiana) responded to inoculation with fungi carried by bark beetles by rapidly increasing monoterpene concentrations at the entry site. Changes in total monoterpenes were more pronounced than changes in proportionate compositions. The extent and rate of host response was affected by fungal species, the viability of the inoculum, and host tree species. In general, host responses were highest to fungi that are phytopathogenic and consistently associated with the major bark beetles in the study region. Simple mechanical wounding cannot account for the observed allelochemical changes, as aseptic inoculations elicited only minor reactions. Similarly, inoculation with autoclaved inviable fungi generally elicited intermediate responses, suggesting that both structural and metabolic fungal properties are important. Responses by jack pine, P. banksiana, were generally more rapid and variable than those of red pine, P. resinosa. Dose-toxicity experiments with synthetic compounds demonstrated that monoterpene concentrations present in vivo only a few days after simulated attack are lethal to most beetles. Constitutive (pre-attack) monoterpene levels can also exert some toxicity. Because bark beetles engage in pheromone-mediated mass attacks that can deplete host defenses, constitutive monoterpene levels, while a necessary early phase of successful plant defense, appear insufficient by themselves. Such interactions between constitutive and induced defense chemistry may be important considerations when evaluating general theories of plant defense.

Gradients of intertidal primary productivity around the coast of South Africa and their relationships with consumer biomass

Abstract: The structure of rocky intertidal communities may be influenced by large-scale patterns of productivity. In this study we examine the in situ rates of production by intertidal epilithic microalgae (chlorophyll a production per unit area per month), intertidal nutrient concentrations (nitrates, nitrites, phosphates and silicates), and standing stocks of different functional-form groups of macroalgae around the South African coast, and their relationships to consumer biomass. Clear gradients of in situ intertidal primary production and nutrient concentrations were recorded around the South African coast, values being highest on the west coast, intermediate on the south and lowest on the east coast. Primary production by intertidal epilithic microalgae was correlated with nutrient availability and could also be related to nearshore phytoplankton production. The dominance patterns of different functional forms of macroalgae changed around the coast, with foliose algae prevalent on the west coast and coralline algae on the east coast. However, overall macroalgal standing stocks did not reflect the productivity gradient, being equally high on the east and west coasts, and low in the south. Positive relationships existed between the average biomass of intertidal intertebrate consumers (grazers and filter-feeders) and intertidal productivity, although only the grazers were directly "connected" to in situ production by epilithic intertidal microalgae. The maximum body size of a widely distributed limpet, Patella granularis, was also positively correlated with level of in situ primary production. The maximal values of biomass attained by intertidal filter-feeders were not related to intertidal primary production, and were relatively constant around the coast. At a local scale, filter-feeder biomass is known to be strongly influenced by wave action. This implies that the local-scale water movements over-ride any effects that large-scale gradients of primary production may have on filter-feeders. The large-scale gradient in intertidal productivity around the coast is thus strongly linked with grazer biomass and individual body size, but any effect it has on filter-feeder biomass seems subsidiary to the local effects of wave action.

Herbivorous mammals as seed dispersers in a Mediterranean dehesa

Abstract: Endozoochrous seed dispersal by herbivorous mammals has been verified repeatedly and its possible influence on the structure and function of herbaceous communities has been suggested. Quantitative studies, however, are lacking in the field of seed dispersal via the dung of herbivore guilds in little-altered environments. The present paper analyses seed dispersal via rabbit, fallow deer, red deer and cow dung in a Mediterranean dehesa (open woodland used for hunting and ranching) during the seeding season. Dung seed content was determined by the glasshouse cultivation of eight dung samples from each herbivore, collected fortnightly between February and August. The four herbivores disperse many seeds (spring averages are 6-15 seeds per gram of dry dung and maxima of 25-70) from a large number of species (totals between 52 and 78). Dispersal seems to be mainly determined by seed production of the plant comminity. This is reflected in (i) the dissemination of a high percentage of the species present in the dehesa, (ii) great seasonal variability, related to seed production, in the amount of seeds and number of species dispersed, and (iii) a high semi-quantitative similarity of seed content in the four types of herbivore dung throughout the year. There is also important quantitative variation that depends on animal traits and feeding habits. These results and the characteristics of species found in dung suggest the adaptation of plant species to the dispersal of their seeds via herbivore gut. This process may well have profound implications for vegetation dynamics and the evolution of plant traits.

Patterns of light and nitrogen distribution in relation to whole canopy carbon gain in C3 and C4 mono- and dicotyledonous species.

Abstract: An analytical model was used to describe the optimal nitrogen distribution From this model, it was hypothesized that the non-uniformity of the nitrogen distribution increases with the canopy extinction rate for light and the total amount of free nitrogen in the canopy, and that it is independent of the slope of the relation between light saturated photosynthesis (Pm) and leaf nitrogen content (nL) These hypotheses were tested experimentally for plants with inherently different architectures and different photosynthetic modes A garden experiment was carried out with a C3 monocot [rice, Oryza sativa (L)], a C3 dicot [soybean, Glycine max (L) Merr] a C4 monocot [sorghum, Sorghum bicolor (L) Moensch] and a C4 dicot [amarantus, Amaranthus cruentus (L)] Leaf photosynthetic characteristics as well as light and nitrogen distribution in the canopies of dense stands of these species were measured The dicot stands were found to have higher extinction coefficients for light than the monocot stands Dicots also had more non-uniform N distribution patterns The main difference between the C3 and C4 species was that the C4 species were found to have a greater slope value of the leaf-level Pm-nL relation Patterns of N distribution were similar in stands of the C3 and C4 species In general, these experimental results were in accordance with the model predictions, in that the pattern of nitrogen allocation in the canopy is mainly determined by the extinction coefficient for light and the total amount of free nitrogen

Impacts of invading N2-fixing Acacia species on patterns of nutrient cycling in two Cape ecosystems: evidence from soil incubation studies and 15N natural abundance values.

Abstract: This study examines the impacts of woody, N2-fixing invasive Acacia spp. on the patterns of nutrient cycling in two invaded ecosystems of differing nutrient status in the Cape floristic region. Patterns of soil nutrient mineralization were measured by a field incubation method while the significance of the fixation process in altering nutrient cycling was assessed by the δ15N natural abundance technique. The results confirm earlier reports that invasion by woody shrubs results in organic matter and nutrient enrichment of surface soils of both ecosystems. However, patterns of nutrient availability (phosphorus and nitrogen) were not necessarily enhanced. In the more fertile strandveld both phosphorus and nitrogen (significant at P<0.10) showed trends towards enhanced annual mineralization rates upon invasion, while in the low nutrient fynbos system only phosphorus followed this trend. It is unclear whether this differential response is a consequence of plant- or soil-derived feedbacks on the decomposition processes in each system. The δ15N values of the soils from the invaded sites of both ecosystems indicated a strong influence of the alien species on the soil nitrogen component. However, as with other studies of natural ecosystems, the contribution of nitrogen from fixation could not be readily quantified with the δ15N natural abundance method because of problems in selecting suitable non-N2-fixing reference plants. A technique of disrupting nodule structure and function, by fumigation with O2, to obtain the δ15N value of a non-N2-fixing speciment of the study species was tried and found to overcome some of the problems associated with the lack of suitable reference plants. With this technique it was possible to detect the almost total dependence of A. saligna on N2-fixation in the fynbos soils with their low nitrogen mineralization rates. In the strandveld ecosystem with much higher soil nitrogen release rates A. cyclops was only partly dependent on fixation (about half) for its nitrogen. The nutrient enrichment of both ecosystems and trends towards enhanced rates of nutrient mineralization could have profound implications on the long-term success of alien invader clearing operations and the restoration of the indigenous flora at these sites.

Woody tissue maintenance respiration of four conifers in contrasting climates.

Abstract: We estimate maintenance respiration for boles of four temperate conifers (ponderosa pine, western hemlock, red pine, and slash pine) from CO2 efflux measurements in autumn, when construction respiration is low or negligible. Maintenance respiration of stems was linearly related to sapwood volume for all species; at 10°C, respiration per unit sapwood volume ranged from 4.8 to 8.3 μmol CO2 m-3 s-1. For all sites combined, respiration increased exponentially with temperature (Q 10 =1.7, r 2=0.78). We estimate that maintenance respiration of aboveground woody tissues of these conifers consumes 52-162 g C m-2 y-1, or 5-13% of net daytime carbon assimilation annually. The fraction of annual net daytime carbon fixation used for stem maintenance respiration increased linearly with the average annual temperature of the site.

Associational resistance and shared doom: effects of epibiosis on herbivory.

Abstract: The potential for spatial associations between palatable and unpalatable plant species to reduce herbivore pressure on the palatable species has been described as associational resistance, associational refuge or associational defense for numerous terrestrial and marine communities. One of the closest associations between species-epibiosis-has not been thoroughly investigated in this regard. In this study we evaluated how different associations between host seaweeds and epibiotic plants and animals influenced the movement of an omnivorous sea urchin (Arbacia punctulata) to the host and subsequent feeding on the host. A. punctulata showed clear preferences when given pairwise choices between 12 prey species (3 animals, 9 algae). These preferences were consistent and allowed us to rank the six epibiont species and six host species linearly from least to most preferred by A. punculata. Most host-epibiont associations dramatically changed urchin preference, increasing or decreasing urchin grazing on fouled hosts as compared to clean conspecifics. Herbivory on the host increased when the epibiont was more preferred, and decreased when it was less preferred than the unfouled host alga. Taking the host species as a point of reference, we classified epibiosis-caused decrease in herbivory as associational resistance, while epibiont-caused increases in herbivory were defined as shared doom. These epibiont-host-herbivore interactions could select for hosts that facilitate the growth of certain low preference epibionts on their surfaces in situations where the resulting decreases in herbivory would offset the various negative effects of being fouled. In contrast, in situations where herbivores are common, the negative effects of being fouled by palatable epibionts may be much greater than is generally assumed. In our assays, unpalatable hosts fouled by palatable epibionts became much more attractive to urchins and rose several ranks on the urchins' preference hierarchy.

Seedling recruitment patterns in a Belizean mangrove forest: effects of establishment ability and physico-chemical factors

Abstract: A field study was conducted to evaluate the relative importance of factors affecting seedling establishment and survival on a mangrove-dominated island in Belize. An examination of spatial patterns of seedling relative densities in relation to reproductive adults and physico-chemical conditions provided correlative information on factors affecting mangrove regeneration patterns. Distance from reproductive adults explained 89–94% of the variation in relative density of Rhizophora mangle seedlings, whereas availability of resources (light and NH4) explained 73–80% of variation in Avicennia germinans seedling relative density. Just after dispersal (December), 89% of the variation in Laguncularia racemosa seedling relative density was attributable to distance from reproductive adults, but 7 months later (July) 74% of the variation was explained by intensity of flooding- and salinity-related stresses. Survivorship (after 2.5 years) of propagules and seedlings of R. mangle and A. germinans transplanted to zones of contrasting physico-chemical conditions demonstrated that: (1) mortality was highest during the establishment phase and major causes were failure to strand before viability was lost, consumption by predators and desiccation; and (2) after establishment, differences in sensitivity to physicochemical stress factors such as flooding (A. germinans) and initial orientation of the seedling axis (R. mangle) exerted a further influence on seedling survival. The results indicate that seedling recruitment in these neotropical forests is strongly influenced by dispersal patterns, differential establishment abilities and effects of physico-chemical factors that vary with elevation and distance from the shoreline.

Environmental and physiological regulation of transpiration in tropical forest gap species: the influence of boundary layer and hydraulic properties.

Abstract: Environmental and physiological regulation of transpiration were examined in several gap-colonizing shrub and tree species during two consecutive dry seasons in a moist, lowland tropical forest on Barro Colorado Island, Panama. Whole plant transpiration, stomatal and total vapor phase (stomatal + boundary layer) conductance, plant water potential and environmental variables were measured concurrently. This allowed control of transpiration (E) to be partitioned quantitatively between stomatal (gs) and boundary layer (gb) conductance and permitted the impact of invividual environmental and physiological variables on stomatal behavior and E to be assessed. Wind speed in treefall gap sites was often below the 0.25 m s−1 stalling speed of the anemometer used and was rarely above 0.5 m s−1, resulting in uniformly low gb (c. 200–300 mmol m−2 s−1) among all species studied regardless of leaf size. Stomatal conductance was typically equal to or somewhat greater than gb. This strongly decoupled E from control by stomata, so that in Miconia argentea a 10% change in gs when gs was near its mean value was predicted to yield only a 2.5% change in E. Porometric estimates of E, obtained as the product of gs and the leaf-bulk air vapor pressure difference (VPD) without taking gb into account, were up to 300% higher than actual E determined from sap flow measurements. Porometry was thus inadequate as a means of assessing the physiological consequences of stomatal behavior in different gap colonizing species. Stomatal responses to humidity strongly limited the increase in E with increasing evaporative demand. Stomata of all species studied appeared to respond to increasing evaporative demand in the same manner when the leaf surface was selected as the reference point for determination of external vapor pressure and when simultaneous variation of light and leaf-air VPD was taken into account. This result suggests that contrasting stomatal responses to similar leaf-bulk air VPD may be governed as much by the external boundary layer as by intrinsic physiological differences among species. Both E and gs initially increased sharply with increasing leaf area-specific total hydraulic conductance of the soil/root/leaf pathway (Gt), becoming asymptotic at higher values of Gt. For both E and gs a unique relationship appeared to describe the response of all species to variations in Gt. The relatively weak correlation observed between gs and midday leaf water potential suggested that stomatal adjustment to variations in water availability coordinated E with water transport efficiency rather than bulk leaf water status.

Predation risk and alternative mating tactics in male Trinidadian guppies (Poecilia reticulata).

Abstract: In the guppy (Poecilia reticulata), males have two alternative mating tactics. Individual males may either display to a receptive female prior to attempting to copulate with her or attempt to quickly sneakcopulate with a female without first displaying to her or without a prior receptive response from her. In this study, I experimentally investigated the effects of simulated local increases in the risk of predation (in the form of a cichlid fish predator model in situ) on the mating tactics used by free-ranging male guppies in two typical macrohabitats (riffle and pool) of a Trinidadian river. Focal male guppies displayed to females significantly less often on average, and conversely attempted sneak copulations more often, in the presence of the predator model than in its absence; this pattern was similar for both habitats. These fish therefore performed a lower proportion of sigmoid displays and increased their sneaky mating attempts when the apparent risk of predation had increased locally. This predator-mediated response is consistent with a trade-off between mating success and risk of mortality due to predation. The results are the first to confirm risk-sensitive mating behaviour in free-ranging male guppies within a population, and demonstrate the potential importance of predators in influencing the relative use of alternative mating tactics in this species on a microgeographical scale in the wild.

Partitioning of water resources among plants of a lowland tropical forest

Abstract: Source water used by plants of several species in a semi-evergreen lowland tropical forest on Barro Colorado Island, Panama, was assessed by comparing the relative abundance of deuterium, D, versus hydrogen, H (stable hydrogen isotope composition, δD) in xylem sap and in soil water at different depths, during the dry season of 1992. Ecological correlates of source water were examined by comparing xylem water δD values with leaf phenology, leaf water status determined with a pressure chamber, and rates of water use determined as mass flow of sap using the stem heat balance method. Soil water δD values decreased sharply to 30 cm, then remained relatively constant with increasing depth. Average δD values were-13‰, for 0–30 cm depth and-36.7‰ for 30–100 cm depth. Soil water δD values were negatively associated with soil water content and soil water potential. Concurrent analyses of xylem water revealed a high degree of partitioning of water resources among species of this tropical forest. Xylem water δD of deciduous trees (average=-25.3±1.4‰) was higher than that of evergreen trees (average=-36.3±3.5‰), indicating that evergreen species had access to the more abundant soil water at greater depth than deciduous species. In evergreen shade-tolerant and high-light requiring shrubs and small trees, δD of xylem water was negatively correlated with transpiration rate and leaf water potential indicating that species using deeper, more abundant water resources had both higher rates of water use and more favorable leaf water status.

Environmental constraints on the growth, photosynthesis and reproductive development of Dryas octopetala at a high Arctic polar semi-desert, Svalbard.

Abstract: Opportunities exist in high Arctic polar semidesert communities for colonisation of unvegetated ground by long-lived clonal plants such as Dryas octopetala. This can be achieved by lateral spread of vegetative ramets, or by sexual reproduction and seedling recruitment. The objectives of this study were (1) to determine whether these two means of proliferation show differential sensitivity to contrasting components of the abiotic environment (temperature, soil nutrient and water availability) and (2) to evaluate the potential for D. octopetala to respond to climate change by an increase in cover and biomass in polar semi-desert communities. Factorial environmental manipulations of growing season temperature, soil nutrient and water status were conducted over 3 years at a polar semi-desert community in Svalbard, Norway (78°56.12′N, 11°50.4′E) and both clonal and sexual reproductive performance, together with instantaneous net photosynthesis (Pn), were recorded during the third season (1993). D. octopetala capitalised rapidly on an amelioration in the availability of inorganic nutrients (N, P and K) by an expansion in leaf area and biomass supported by increased Pn per unit leaf weight, and by apparent luxury uptake of nutrients (particularly P). Several facets of sexual reproductive development and seed viability were markedly improved by elevated temperatures or soil nutrient availability. Thus although D. octopetala is a long-lived clonal plant, with many traits characteristic of stress resistance syndrome, it showed considerable phenotypic plasticity in response to environmental manipulations. The results support the hypothesis that clonal growth confers survival potential during unfavourable years, together with the ability to capitalise on nutrient flushes and recycle nutrients internally. Continued investment in sexual reproduction ensures that seed setting is successful during favourable years, even if these occur infrequently. Cimate warming in the high Arctic could thus result in changes in the cover, biomass and the relative significance of clonal versus sexual proliferation of D. octopetala (and thus the genetic diversity of the population) but the long-term responses will probably be constrained by lack of available nutrients.

Adaptation of ruminants to browse and grass diets: are anatomical-based browser-grazer interpretations valid?

Abstract: As a result of pioneering work of Hofmann (1973, 1989), nutritional ecologists classify ruminants into three feeding-type categories: browsers (“concentrate” feeders), grazers, and intermediate or mixed feeders. Hofmann proposed that these feeding types result from evolutionary adaptations in the anatomy of the digestive system and that one consequence is shorter retention of the digesta in the rumen of browsers, and thus a decreased efficiency of fiber digestion relative to that of grazers. We examined the hypotheses that (1) fiber digestion of browsers is lower than that of grazers, (2) salivary gland size is larger in all browsers than in grazers, (3) the browser's larger salivary glands produce larger volumes of thin serous saliva than those of grazers, and (4) thus, browsers have higher liquid passage rates than do grazers. We found that the extent of fiber digestion is not significantly different between browsers and grazers, although fiber digestion is positively related to herbivore size. In general, salivary gland size is approximately 4 times larger in browsers than grazers, but some browsers (e.g., greater kudu) have small, grazer-sized salivary glands. Resting (non-feeding or ruminating) saliva flow rates of mule deer (browser) and domestic sheep and cattle (grazers) were not significantly different from each other. Finally, ruminal liquid flow rates were not different between feeding types. We conclude that many of Hofmann's nutritional and physiological interpretations of anatomical differences amongst ruminants are not supportable.

Intraspecific variation in aspen phytochemistry: effects on performance of gypsy moths and forest tent caterpillars

Abstract: Individual quaking aspen trees vary greatly in foliar chemistry and susceptibility to defoliation by gypsy moths and forest tent caterpillars. To relate performance of these insects to differences in foliar chemistry, we reared larvac from egg hatch to pupation on leaves from different aspen trees and analyzed leaf samples for water, nitrogen, total nonstructural carbohydrates, phenolic glycosides, and condensed tannins. Larval performance varied markedly among trees. Pupal weights of both species were strongly and inversely related to phenolic glycoside concentrations. In addition, gypsy moth performance was positively related to condensed tannin concentrations, whereas forest tent caterpillar pupal weights were positively associated with leaf nitrogen concentrations. A subsequent study with larvae fed aspen leaves supplemented with the phenolic glycoside tremulacin confirmed that the compound reduces larval performance. Larvae exhibited increased stadium durations and decreased relative growth rates and food conversion efficiencies as dietary levels of tremulacin increased. Differences in performance were more pronounced for gypsy moths than for forest tent caterpillars. These results suggest that intraspecific variation in defensive chemistry may strongly mediate interactions between aspen, gypsy moths and forest tent caterpillars in the Great Lakes region, and may account for differential defoliation of aspen by these two insect species.

Leaf dynamics, self-shading and carbon gain in seedlings of a tropical pioneer tree.

Abstract: We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.

Carbon and nitrogen mineralization from decomposing gypsy moth frass.

Abstract: Defoliation of forests by insects is often assumed to produce a pulse of available nitrogen (N) from the decomposition of frass pellets. In this study we measured rates of carbon (C) and N mineralization from gypsy moth frass incubated with and without soil, and for soil alone. Incubations were at constant temperature and soil moisture conditions and lasted for 120 days. We found that gypsy moth frass contains much labile C as well as extractable N, and that the stimulation of microbial growth by the labile C results in immobilization of essentially all of the extractable N in the frass. The response of the microbes is fast, beginning within 1 day and lasting at least 90 days. This immobilization response represents an efficient mechanism for conserving N within a forest ecosystem after a defoliation event.

Dietary energetics of the insectivorous Mexican free-tailed bat (Tadarida brasiliensis) during pregnancy and lactation.

Abstract: Stomach content analysis of 20 pregnant (x body mass=13.4 g) and 18 lactating (x body mass=11.5 g) female Tadarida brasiliensis revealed that the diet, expressed as percent volume, consists largely of lepidopterans, coleopterans, hymenopterans, and dipterans, in decreasing order of importance. We found no significant difference in the diet of pregnant and lactating females when expressed as percent volume. However, when expressed as percent frequency, proportionately more pregnant females fed on lepidopterans, coleopterans, and dipterans than did lactating bats, and proportionately more lactating females fed on hymenopterans. We found no significant differences in the percentages of water, lean dry mass, fat, and energy density in the stomach contents of pregnant and lactating females. Water in stomach contents averaged 62.7% and fat and lean dry mass averaged 22.2% and 15.2%, respectively (expressed as percentage of wet mass); energy density averaged 31.2 kJ g−1 dry mass. This relatively high energy density of stomach contents, as compared to whole insects, can be attributed to the consumption of insects high in fat (especially flying ants) and the abdomens only of moths and beetles (other body parts being discarded). Estimates of nightly food intake increased markedly from mid- to late pregnancy, stabilized or decreased during late pregnancy, and increased again during early to mid-lactation. Average nightly feeding rate doubled from pregnancy to lactation and increased threefold during the first half of lactation. By adjusting our gravimetric estimates of nightly feeding rate upwards by 40% (based on estimates of field metabolic rate), we estimate that the average nightly energy intake of female T. brasiliensis ranges from 57 kJ day−1 in early lactation to 104 kJ day−1 in mid-lactation. These estimates represent nightly feeding rates averaging 39% and 73% of a female's body mass in the period from early to mid-lactation, respectively.

Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna.

Abstract: Most savanna water balance models assume water partitioning between grasses and shrubs in a two-layer hypothesis, but this hypothesis has not been tested for humid savanna environments. Spatial partitioning of soil water between grasses and shrubs was investigated in a West African humid savanna by comparing the isotopic composition (oxygen-18 and deuterium) of soil water and plant stem water during rainy and dry conditions. Both grass and shrub species acquire most of their water from the top soil layer during both rainy and dry periods. A shift of water uptake pattern towards deeper horizons was observed only at the end of the dry season after shrub defoliation. The mean depth of water uptake, as determined by the isotopic signature of stem water, was consistent with grass and shrub root profiles and with changes in soil water content profiles as surveyed by a neutron probe. This provides evidence for potentially strong competition between shrubs and grasses for soil water in these humid savannas. Limited nutrient availability may explain these competitive interactions. These results enhance our understanding of shrub-grass interactions, and will contribute to models of ecosystem functioning in humid savannas.

Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment.

Abstract: Acacia karroo Hayne is the most important woody invader of grassland in South Africa, and can greatly reduce the productivity of grassland A field experiment was conducted to test the hypotheses that emergence, growth and the 1st year's survival of Acacia karroo would be enhanced by (1) defoliation of the grass sward, (2) increased irradiance, (3) increased moisture availability and (4) its germination within cattle dung pats The study was conducted on one site above and one below the natural altitudinal treeline of this species in grassland of the eastern Cape, South Africa Not one seedling emerged from dung pats Neither location nor the other treatments affected the density of emerging seedlings, although only 404 seedlings m−2 emerged of the 200 seeds m−2 planted Shading dramatically increased the density of surviving seedlings In the open, only 3 and 15 seedlings m−2 remained respectively at the end of the growing season or the beginning of the next, compared to 233 and 195 seedlings m−2 under shading for these respective times This was attributed to the effect of shade on moisture availability in a season which received only 54% of average rainfall Seedling survival until the end of the growing season was enhanced (30%) by shade at both sites, but also by supplemental water (24%) and defoliation of the sward (7%) at the site above the treeline Across sites and treatments, seedling survival was related to moisture availability, with no or poor survival for < 500 mm rainfall, indicating this species can only establish in certain rainfall years Seedling survival over winter was not influenced by treatment, but was greater for larger seedlings Treatments affected seedling size, in particular seedlings growing under shade and within a dense grass canopy were etiolated A karroo seedlings are capable of establishing and surviving within a dense grass sward for at least a year, tolerant of low irradiance and of interference, which, because most seeds do not persist for much longer than a year, suggests this species forms predominantly a seedling bank This has implications for the invasion of grassland by woody species

Estimation of deciduous forest leaf area index using direct and indirect methods

Abstract: This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the “Demon” portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6–37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI−WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.

Leaf and canopy responses to elevated CO2 in a pine forest under free-air CO2 enrichment.

Abstract: Physiological responses to elevated CO2 at the leaf and canopy-level were studied in an intact pine (Pinus taeda) forest ecosystem exposed to elevated CO2 using a free-air CO2 enrichment (FACE) technique. Normalized canopy water-use of trees exposed to elevated CO2 over an 8-day exposure period was similar to that of trees exposed to current ambient CO2 under sunny conditions. During a portion of the exposure period when sky conditions were cloudy, CO2-exposed trees showed minor (≤7%) but significant reductions in relative sap flux density compared to trees under ambient CO2 conditions. Short-term (minutes) direct stomatal responses to elevated CO2 were also relatively weak (≈5% reduction in stomatal aperture in response to high CO2 concentrations). We observed no evidence of adjustment in stomatal conductance in foliage grown under elevated CO2 for nearly 80 days compared to foliage grown under current ambient CO2, so intrinsic leaf water-use efficiency at elevated CO2 was enhanced primarily by direct responses of photosynthesis to CO2. We did not detect statistical differences in parameters from photosynthetic responses to intercellular CO2 (A net-C i curves) for Pinus taeda foliage grown under elevated CO2 (550 μmol mol−1) for 50–80 days compared to those for foliage grown under current ambient CO2 from similar-sized reference trees nearby. In both cases, leaf net photosynthetic rate at 550 μmol mol−1 CO2 was enhanced by approximately 65% compared to the rate at ambient CO2 (350 μmol mol−1). A similar level of enhancement under elevated CO2 was observed for daily photosynthesis under field conditions on a sunny day. While enhancement of photosynthesis by elevated CO2 during the study period appears to be primarily attributable to direct photosynthetic responses to CO2 in the pine forest, longer-term CO2 responses and feedbacks remain to be evaluated.

The influence of colonization in nested species subsets.

Abstract: Biotic communities inhabiting collections of insular habitat patches often exhibit compositional patterns described as “nested subsets”. In nested biotas, the assemblages of species in relatively depauperate sites comprise successive subsets of species in relatively richer sites. In theory, nestedness may result from selective extinction, selective colonization, or other mechanisms, such as nested habitats. Allopatric speciation is expected to reduce nestedness. Previous studies, based largely on comparisons between land-bridge and oceanic archipelagos, have emphasized the role of selective extinction. However, colonization could also be important in generating strong patterns of nestedness. We apply a recently published index of nestedness to more than 50 island biogeographic data sets, and examine the roles of colonization, extinction, endemism, and, to a limited extent, habitat variability on the degree on nestedness. Most data sets exhibit a significant degree of nestedness, although there is no general tendency for land-bridge biotas to appear more nested than oceanic ones. Endemic species are shown to generally reduce nestedness. Comparisons between groups of non-endemic species differing in overwater or inter-patch dispersal ability indicate that superior dispersers generally exhibit a greater degree of nestedness than poorer dispersers, a result opposite that expected if colonization were a less predictable process than extinction. These results suggest that frequent colonization is likely to enhance nestedness, thereby increasing the compositional overlap among insular biotas. The prevalence of selective extinction in natural communities remains in question. The importance of colonization in generating and maintaining nested subsets suggests that (1) minimum critical areas will be difficult to determine from patterns of species distributions on islands; (2) multiple conservation sites are likely to be required to preserve communities in subdivided landscapes; and (3) management of dispersal processes may be as important to preserving species and communities as is minimizing extinctions.

Slow larval growth on a suboptimal willow results in high predation mortality in the leaf beetle Galerucella lineola.

Abstract: The slow growth/high mortality hypothesis predicts that herbivorous insects feeding on suboptimal host plants are subjected to higher predation mortality owing to the longer time spent in the vulnerable juvenile stages compared with conspecifics feeding on optimal plants. We tested this hypothesis for the willow-feeding leaf beetle Galerucella lineola raised on one suitable (Salix viminalis) and one unsuitable (S. dasyclados) willow species as well as on plants from an interspecific cross between the two species. Cohorts of larvae raised on caged plants (protected from enemies) and uncaged plants (exposed to enemies) were monitored daily throughout larval development in two consecutive years. Larvae raised on S. viminalis developed faster, grew larger and survived better than those raised on S. dasyclados. The suitability of the hybrid plants was intermediate to that of their parents. Our results strongly support the slow growth/high mortality hypothesis. In both years, total predation during the larval period was higher on S. dasyclados than on S. viminalis. Furthermore, the daily predation rate (i.e. the proportion of larvae preyed upon per day) was higher on S. dasyclados than on S. viminalis. When hybrid plants were included in the analysis total predation was positively correlated with both larval development time and daily predation rate. We suggest that high predation on beetles on low-quality plants is the combined result of their longer development time and elevated daily predation rate. The results are discussed in relation to the evolution of host plant selection and the paradox of sublethal plant defenses.

The fate of 15N-labelled nitrate additions to a northern hardwood forest in eastern Maine, USA.

Abstract: We followed the movements of 15N-labelled nitrate additions into biomass and soil pools of experimental plots (15×15 m each) in a mid-successional beech-maple-birch-spruce forest in order to identify sinks for nitrate inputs to a forest ecosystem Replicate plots (n=3) were spray-irrigated with either 28 or 56 kg N ha-1 year-1 using 15N-labelled nitric acid solutions (δ15N = 344‰ ) during four successive growing seasons (April-October) The 15N contents of foliage, bolewood, forests floor and mineral soil (0-5 cm) increased during the course of treatments Mass balance calculations showed that one-fourth to one-third of the nitrate applied to forest plots was assimilated into and retained by above ground plant tissues and surface soil horizons at both rates of nitrate application Plant and microbial assimilation were of approximately equal importance in retaining nitrate additions to this forest Nitrate use among tree species varied, however, with red spruce showing lower rates of nitrate assimilation into foliage and bolewood than American beech and other deciduous species

The effect of matrix on the occurrence of hazel grouse (Bonasa bonasia) in isolated habitat fragments.

Abstract: The aim of this study was to determine the effect of matrix on the occurrence of hazel grouse (Bonasa bonasia) in habitat fragments. The study was conducted in two kinds of landscape: (1) an agricultural landscape, where the censused forest habitat fragments were surrounded by farmland, and (2) in an intensively managed forested landscape, where the censused habitat fragments were surrounded by nonhabitat coniferous forest. Occupied and unoccupied habitat fragments in the agricultural landscape differed significantly in distance to the nearest suitable continuous habitat, with hazel grouse occurring only in habitat fragments closer than 100 m from continuous forest. In the intensively managed forest landscape, the effect of isolation was less evident, but there might be a threshold around 2 km. Effects of isolation occurred over much shorter distances when the surrounding habitats consisted of farmland than when it was forested habitats. The size of the habitat fragments was important in both landscapes, with larger habitat fragments more often containing hazel grouse.