Demographic responses of Pinguicula ionantha to prescribed fire: a regression-design LTRE approach

This study describes the use of periodic matrix analysis and regression-design life table response experiments (LTRE) to investigate the effects of prescribed fire on demographic responses of Pinguicula ionantha, a federally listed plant endemic to the herb bog/savanna community in north Florida. Multi-state mark–recapture models with dead recoveries were used to estimate survival and transition probabilities for over 2,300 individuals in 12 populations of P. ionantha. These estimates were applied to parameterize matrix models used in further analyses. P. ionantha demographics were found to be strongly dependent on prescribed fire events. Periodic matrix models were used to evaluate season of burn (either growing or dormant season) for fire return intervals ranging from 1 to 20 years. Annual growing and biannual dormant season fires maximized population growth rates for this species. A regression design LTRE was used to evaluate the effect of number of days since last fire on population growth. Maximum population growth rates calculated using standard asymptotic analysis were realized shortly following a burn event (<2 years), and a regression design LTRE showed that short-term fire-mediated changes in vital rates translated into observed increases in population growth. The LTRE identified fecundity and individual growth as contributing most to increases in post-fire population growth. Our analyses found that the current four-year prescribed fire return intervals used at the study sites can be significantly shortened to increase the population growth rates of this rare species. Understanding the role of fire frequency and season in creating and maintaining appropriate habitat for this species may aid in the conservation of this and other rare herb bog/savanna inhabitants. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effects of pollen limitation on population dynamics of snow lotus (<Emphasis Type="Italic">Saussurea medusa</Emphasis> and <Emphasis Type="Italic">S. laniceps</Emphasis>, Asteraceae): Threatened Tibetan medicinal plants of the eastern Himalayas

Pollen limitation reduces seed production and may reduce plant population growth rate. Plants may be particularly prone to pollen limitation if they require pollinators, occur at high elevation, and have human-mediated reductions in plant density due to harvesting. We found that two rare monocarpic Tibetan plant species, known as the Himalayan Snow Lotus, both require pollinators and that seed production in Saussurea medusa (Asteraceae) but not S. laniceps is limited by pollen receipt. We created deterministic and stochastic stage-structured matrix models for S. medusa, and found that pollen supplementation significantly increases population growth rate. However, even when pollen is not limiting, S. medusa is likely at risk for extinction in the next 50 years. Our results for this monocarpic plant differ from other population studies on pollen limitation in polycarpic plant species since the magnitude of pollen limitation for seed production was relatively low, and yet the sensitivity of population growth to changes in seed production was relatively high.     

Sampling Bias in Roadsides: The Case of Galling Aphids on Pistacia Trees

Sampling along roadsides is convenient and is widely practiced in insect population researches. Ecological conditions in road verges are very different than those prevailing in natural habitats and they affect the annual growth of plants in semi-arid and arid regions. This in turn may improve development, survival and abundance of insects feeding on plants growing in roadsides. These trends may bias the results of sampling. To verify this assertion, we quantified the effects of growing in roadside on annual growth of Pistacia atlantica trees and Pistacia palaestina shrubs and compare two demographic indexes of nine gall-inducing aphid species on trees growing along roads with trees in the open landscape, in Israel. The annual growth of the two host plants was significantly more vigorous in roadsides than away from roads. Tests of Combined Probabilities showed that the likelihood of P. atlantica and P. palaestina to be parasitized by more galls of Fordini species is higher in roadsides than away from roads. Moreover, in the semi-dry regions of Israel, three aphid species on P. atlantica and five species on P. palaestina induced more galls in plants growing along roads than away from roads, while in the rainy Northern region, the difference was not significant between the two habitats. These results indicate a biased evaluation of population size in roadside habitat, which has to be accounted in insect–plant relation researches.     

Comparative demography of two giant caulescent rosettes (Espeletia timotensis and E. spicata) from the high tropical Andes

Using field data from previous studies we built matrix models for two populations of giant rosettes, Espeletia timotensis Cuatrec. and E. spicata Sch. Bip. Wedd., from the Andes Cordillera in Mérida, Venezuela. We analysed the models and calculated population growth rate (λ), sensitivities, elasticities and the sensitivity of the elasticities to changes in the vital rates. The analysis showed that the two species behave alike in general demographic terms. In both models, population growth rate is positive and sensitivities of λ to changes in vital rates decrease markedly in this order: plant establishment, progression of juvenile–adult, germination and survival. The relative contributions of vital rates to λ (elasticities) are very similar to those of other woody plant species: a higher contribution of survival and a very low contribution of fecundity. Transition from seedling to juvenile is most important and the younger established stages (juveniles and young adults) play a predominant demographic role in both populations. Seed banks and older adults are playing a relatively minor role in the dynamics of both populations. However, they may be important in relation to unpredictable, favourable or detrimental events. Perturbation analysis of elasticities showed that increasing the rate of plant establishment will decrease the relative importance of stasis. We conclude that both species are demographically very close, and similar to other long‐lived woody plant species. However, the two species differ in the role of the seed bank, which seems more important in the demography of E. spicata than in E. timotensis.     

Reference data for evaluating the growth of common riverine fishes in the UK

Comparative assessments of population mean growth rates in length remain important aspects of stock assessment in river fisheries. To facilitate these assessments, for 15 fish species encountered in UK rivers reference data are provided on their expected lengths at age, maximum theoretical lengths (L_∞), growth coefficient (K) and instantaneous mortality rate (Z). These data are also transferable to fish populations outside of the UK that experience a similar growth season (approximately April to October, mean water temperatures 12–22°C). Considerable plasticity was observed in the growth of all species examined, with length at age values revealing growth rates from very slow to very fast. Populations considered fast growing against reference data were coincident with a relatively low ultimate length, a high growth coefficient and a high instantaneous mortality rate, suggesting a trade‐off exists between growth rate, ultimate length and life span.     

Life history patterns of parental and hybrid Daphnia differ between lakes

1. We investigated whether Daphnia galeata × hyalina hybrids of Lake Constance and Lake Greifensee show the same pattern of life history parameters as previously reported for D. galeata × cucullata hybrids and whether such a pattern is consistent between Daphnia populations from those two lakes. 2. Hybrids in Lake Constance were intermediate in size compared with the parental species. Hybrids in Lake Greifensee were smaller than D. galeata. The intrinsic growth rate (r) of hybrids from Lake Constance was not significantly different from the faster growing parental taxon D. galeata. However, r of hybrids from Lake Greifensee was significantly lower than that of D. galeata. 3. The observed juvenile body length differences between the taxa varied with the clutch number. The first clutch juvenile lengths of the three taxa did not differ for Lake Constance. First clutch juveniles of Lake Greifensee D. galeata were smaller than hybrid first clutch juveniles. The third clutch juvenile length did not differ between taxa from Lake Greifensee, but D. galeata juveniles from Lake Constance were bigger than those of D. hyalina. 4. The life history pattern found in Lake Constance corresponds to previous findings from other studies. The hybrids in this lake combine the faster population growth of one parental species with a relatively small size. In the case of Lake Greifensee hybrids, the relatively large size of first clutch juveniles and the small size of the adults could be interpreted as dual adaptations to invertebrate and fish predation. We speculate that the lower population growth rate of the hybrids is a trade‐off for this twofold protection.     

Simulation of the effects of Acanthaster planci on the population structure of massive corals in the genus Porites: evidence of population resilience?

Scleractinian corals in the genus Porites are slow growing, can live for centuries, and can attain great size. In these respects they differ from the majority of coral species, which grow faster and live for years to decades. The predatory starfish Acanthaster planci L. feeds on a wide range of coral species including Porites spp., and during outbreaks in its populations, causes high coral mortality and injury over much of the affected reefs. Because they are slow growing and because recent outbreaks of the starfish occurred only 15 years apart, it may be argued that the Porites populations on affected reefs will be sent into a period of prolonged decline. The present study uses a size stage model of the transition matrix type to predict effects of starfish outbreaks of various frequencies on Porites populations. A transition matrix characterizing the mortality and injury caused in different Porites size classes at John Brewer Reef during an outbreak year was determined from field data. Transition matrices for non-outbreak years were constructed on the basis of realistic growth rates and postulated survivorship and recruitment schedules. The medium term (100 years) effects of outbreaks were simulated by alternation of a single iteration of the outbreak matrix with many iterations of each non-outbreak matrix. By varying the interval between simulated outbreaks it was possible to define combinations of growth rate, survivorship and recruitment which were viable for various outbreak intervals. Simulations based on estimates of the initial size frequency distribution, recruitment rates and colony growth rates for the John Brewer Reef population predicted that the population would remain viable in the face of outbreaks every 15 years only if juvenile and adult survivorship were high. However, within the range of colony growth rates known to occur throughout the Great Barrier Reef and at recruitment rates of the same order as those estimated in the field population, it appears that a much wider range of survivorship schedules could lead to parity or even sustained growth in the face of outbreaks recurring at intervals of from 1 to 3 decades. It is suggested that because the key measurable parameters (initial size structure, damage characteristics, recruitment rate and growth rate) are likely to be very patchy at the scale of whole reefs, no general statement concerning the prognosis for Porites would be meaningful. However the model provides a tool by which a standardized evaluation of this type of field data may be made on a reef by reef basis.     

Conservation prospects for threatened Vietnamese tree species: results from a demographic study

Given that changes in population size are slow, information on future prospects of long-lived tree species is necessarily obtained from demographic models. We studied six threatened tree species in four Vietnamese protected areas: the broad-leaved Annamocarya sinensis, Manglietia fordiana and Parashorea chinensis, and the coniferous Calocedrus macrolepis, Dacrydium elatum and Pinus kwangtungensis. With data from a 2-year field study on recruitment, growth and survival, we constructed matrix models for each species. All species showed continuous regeneration, as indicated by annual seedling recruitment and inverse J-shaped population structures. To evaluate the future prospects of our study species, we calculated three parameters: (1) asymptotic growth rates (λ) from matrix models indicated significant population declines of 2–3%/year for two species; (2) population trajectories for 50–100 years showed slight population declines (0–3%/year) for five species; and (3) the reproductive period required for an adult tree to replace itself was excessive for three of the six species, suggesting that these species presently have insufficient recruitment. Overall agreement of the three parameters was low, showing that reliance on just one parameter is risky. Combining the three parameters we concluded that prospects are good for Dacrydium and Parashorea, worrisome for Annamocarya, Manglietia and Pinus, and intermediate for Calocedrus. We argue that conservation should involve strict protection of (pre-)adult trees, as their survival is crucial for population maintenance in all species (high elasticity). For species with poor demographic prospects, active intervention is required to improve seedling and tree growth, enrich populations with seedlings from controlled germination, and restore habitat. Finally, our study suggests that these conservation measures apply to long-lived trees in general, given that their demography is highly similar. Such measures should be taken before populations decline below critical levels, as long-lived species will respond slowly to management. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Comparative demographic analysis in contrasting environments of Magnolia dealbata: an endangered species from Mexico

The study of population regulation and demography in natural habitats is critical for the conservation of rare and endangered species. We address the impact of cattle exclusion on the demographics of Magnolia dealbata (Magnoliaceae), an endangered species according to IUCN and Federal Mexican laws. Sixteen permanent plots were established, eight of which were enclosed to livestock, in the mountain cloud forest in Coyopolan, Mexico, which has the largest population of M. deadlbata. Censuses of the plots were undertaken annually during three annual cycles to record seed number, recruitment, mortality, and growth (defined as length and diameter at breast height). The effects of two treatments (with and without livestock exclusion) on the dynamics of M. dealbata were investigated using transition matrix models and life table response experiment (LTRE) analysis. Contrary to expectations, there was no significant effect of cattle exclusion on population growth rates (λ). Furthermore, the λ, estimated from the mean transition matrix for both treatments was greater than one. The transitions with the highest elasticity values were similar between the two treatments, while reproductive stage contributed more to differences in population growth rate and were less variable than the non-reproductive stage. LTRE analysis showed that treatment differences had little effect on λ. Livestock exclusion appears to lead more to differences in the arrangement of the values of the transition matrix than to the rate of population growth for M. dealbata.     

Changes in <Emphasis Type="Italic">Synechococcus</Emphasis> Population Size and Cellular Ribosomal RNA Content in Response to Predation and Nutrient Limitation

A mathematical model of predator-prey interactions was used to predict the relationship between population size and cellular growth rate in a two-tiered trophic system consisting of Synechococcus PCC 6301 and Tetrahymena pyriformis. As predicted, axenic chemostat cultures of Synechococcus responded to increased nutrient availability by expanding the equilibrium population size without a concurrent change in growth rate. Likewise, the addition of the predator Tetrahymena pyriformis decreased the Synechococcus population size by 85% and increased the Synechococcus growth rate. Synechococcus populations in the surface waters of the Gulf of Mexico were sampled to ascertain whether the relationship between population size and cellular 16S rRNA concentration conformed to that predicted by the model. Direct counts of autofluorescent cells in size-fractionated seawater samples provided an estimate of Synechococcus population size. The growth rate of in situ populations was estimated by measuring the extent of hybridization of an oligonucleotide probes complementary to Synechococcus 16S rRNA, based on evidence that ribosomal RNA content increases concurrently with growth rate. The comparison of in situ population sizes and specific growth rates revealed that relatively large Synechococcus populations were growing slowly, indicative of nutrient limitation, and that quickly growing populations were relatively small, as predicted for predator-limited populations.     

Population dynamics and comparative demographics in sympatric populations of the round-leaved orchids Platanthera macrophylla and P. orbiculata

Orchids (Orchidaceae) are a family of flowering plants with a high proportion of threatened taxa making them an important focus of plant conservation. Orchid conservation efforts are most effective when informed by reliable demographic research. We utilized transition matrix models to examine the population dynamics and demography within sympatric populations of a species pair of terrestrial round-leaved orchids, Platanthera macrophylla and P. orbiculata. The models were parameterized from a large data set spanning 9 years from field observations of over 1,000 orchids. Life table response experiments (LTRE) were used to identify which life history transitions, and which vital rates within those transitions, most contributed to observed differences between the two species and most contributed to interannual variation within each species. Results from mean transition matrices projected finite rates of population growth that were not significantly different between the two species, with P. macrophylla near the replacement rate and P. orbiculata below it. LTRE revealed that the difference in population growth rates between the two species was mostly due to differences in fecundity (flowering adult to protocorm transition) driven by differences in fruit set and seed germination into protocorm, which were much greater for P. macrophylla. The two primary contributors to interannual variation in population growth rates for both orchids were adult survival and fruit set, respectively. These findings indicate that any environmental disturbances harming adult survival or fecundity will have a disproportionately negative effect on the orchid populations.     

Effects of canopy gaps on the demography of the subcanopy tree Styrax obassia

Abstract. To evaluate the gap dependency of the subcanopy tree species Styrax obassia, we estimated the demographic parameters of this species in a permanent plot in the Ogawa Forest Reserve, Central Japan, a temperate deciduous forest. From the data collected over eight years, we constructed four transition matrix models of Styrax: a whole population; a shaded subpopulation; a gap-site subpopulation; a compound population of shaded and gap-site subpopulations. The whole-population model suggested that the population structure of Styrax is stable in its present condition. The asymptotic population growth rate of the shaded subpopulation was larger than 1, suggesting that this species can maintain its population even under shaded conditions. The elasticity analyses showed the relative importance of survival of adult individuals that stay mostly under a closed canopy. Since the subpopulation in gaps is very small in a mature forest, the contributions of high seedling survival and high fecundity of adult trees in gaps to the whole population growth were smaller than the survival of adult individuals in the shade. Although we showed that this species has the potential to increase its population in larger disturbances, it actually dominates in the study site where the disturbance scale is relatively small. Styrax should be classified as a typical shade-tolerant species, and it also takes good advantage of small scale disturbance. It can sufficiently maintain its population in a mature forest with small-scale disturbances and has the adaptive characteristics of species that live under a closed canopy.     

Using integral projection models to compare population dynamics of four closely related species

Demographic processes, such as survival, growth, and reproduction, can inform us about invasion risk, extinction risk, and trade-offs in life history strategies. The population dynamics of four Amaranthaceae species in southern Illinois, USA were examined using integral projection models (IPMs) to determine whether vital rates reflect life history among these closely related species. Two of the species, Amaranthus palmeri and Amaranthus tuberculatus, are summer annuals and considered to be some of the most problematic agricultural weeds in the US Midwest. Achyranthes japonica is a relatively new invasive exotic species that primarily inhabits forests. Iresine rhizomatosa, is an endangered species in the study area, which also inhabits forests. Two populations of each species were studied from 2012 to 2014 in which height of individuals were measured and used as the state variable in the IPMs. The Amaranthus species and Achyranthes japonica had an estimated population growth rate >1, projecting increases in population size. By contrast, λ was <1 for I. rhizomatosa, projecting a decline in population size demonstrating its endangered status. Germination rates and seed viability were dependent on species and varied over time. Elasticity analyses showed that survival and growth contributed most to λ for the perennial species; whereas, for the annual species population dynamics were driven primarily by fecundity. Overall, Achyranthes japonica and the Amaranthus species show similar trends in demographic processes that align with their invasive nature and not with their life histories. Furthermore, this study demonstrates that more research on the competitive nature of Achyranthes japonica is needed.     

Calculating growth parameters of Genidens barbus (Siluriformes, Ariidae) using length composition and age data

The marine catfish Genidens barbus was once one of the most important estuarine fishery resources in the Patos Lagoon (southern Brazil), which had the largest population known for the species. However, clear signs of overfishing have been observed since the 1980s. Growth parameters for this population were estimated by combining length frequency data for juveniles and selected published data on annuli reading from adult otoliths. This methodology yielded a more realistic set of parameters using both the specialized and generalized von Bertalanffy growth model. The species growth can be described as: L_t = 118.6[1 − e^{−0.043(t+1.505)}] or L_t = 104.6[1 −e^{−0.137D(t+4.013)}]^1/D with the surface factor D = 0.542. The growth performance index ϕ′ was estimated as 2.78 for the special (classical) model. Stock rebuilding is expected to be slow, as the species is slow‐growing, has a low natural mortality rate, and mouthbreeds a small amount of large eggs.     

Life history of the red-banded seabream Pagrus auriga (Sparidae) from the coasts of the Canarian archipelago

Red‐banded seabream Pagrus auriga (N = 615) were caught off the Canary Islands from January 2003 to December 2004. Total length ranged from 120 to 780 mm. The species was characterized by protogynous hermaphroditism. The male :female ratio was in favour of females (1 : 8.2). The reproductive season extended from September to February, with a peak in spawning activity in October–November. Fifty percentage maturity was reached at 387 mm total length by females and 533 mm by males. The length–weight relationship for all individuals was described by the parameters: a = 0.0086 and b = 3.014, when length is given in mm and weight in grams. Otolith age readings indicated that the population consists of 19 age groups, including a very high proportion of individuals between 0 and 7 years old. Growth analysis reveals that the species is slow‐growing and relatively long lived (18 years). The von Bertalanffy growth parameters for the entire population were: L_∞ = 803 mm, k = 0.081 year⁻¹ and t₀ = −2.17 year. Growth differed between males and females. The instantaneous rate of natural mortality for all fish was: M = 0.164 year⁻¹.     

A modelling approach to estimate the effect of exotic pollinators on exotic weed population dynamics: bumblebees and broom in Australia

The role of mutualisms in contributing to species invasions is rarely considered, inhibiting effective risk analysis and management options. Potential ecological consequences of invasion of non‐native pollinators include increased pollination and seed set of invasive plants, with subsequent impacts on population growth rates and rates of spread. We outline a quantitative approach for evaluating the impact of a proposed introduction of an invasive pollinator on existing weed population dynamics and demonstrate the use of this approach on a relatively data‐rich case study: the impacts on Cytisus scoparius (Scotch broom) from proposed introduction of Bombus terrestris. Three models have been used to assess population growth (matrix model), spread speed (integrodifference equation), and equilibrium occupancy (lattice model) for C. scoparius. We use available demographic data for an Australian population to parameterize two of these models. Increased seed set due to more efficient pollination resulted in a higher population growth rate in the density‐independent matrix model, whereas simulations of enhanced pollination scenarios had a negligible effect on equilibrium weed occupancy in the lattice model. This is attributed to strong microsite limitation of recruitment in invasive C. scoparius populations observed in Australia and incorporated in the lattice model. A lack of information regarding secondary ant dispersal of C. scoparius prevents us from parameterizing the integrodifference equation model for Australia, but studies of invasive populations in California suggest that spread speed will also increase with higher seed set. For microsite‐limited C. scoparius populations, increased seed set has minimal effects on equilibrium site occupancy. However, for density‐independent rapidly invading populations, increased seed set is likely to lead to higher growth rates and spread speeds. The impacts of introduced pollinators on native flora and fauna and the potential for promoting range expansion in pollinator‐limited ‘sleeper weeds’ also remain substantial risks.     

Population responses of Ulex shrubs to fire in a lowland heath community

Question: How does the frequency of heathland fire events affect population growth rates of two woody shrub species, Ulex gallii and U. minor? Location: Dry heathland on the south coast of England, UK. Methods: The population dynamics of U. gallii and U. minor were modelled at each phase of the heathland cycle — pioneer, building, mature and degenerate — using periodic matrix products to investigate the response to different fire regimes. Results: Population growth rates of both Ulex species declined under annual burning. Initially, as the time between burns increased, population growth rate increased for both species. Maximum population growth rates for each Ulex species were achieved under a 16‐yr fire return interval. Fire return intervals > 16 yr resulted in declining population growth rates. Conclusions: A species‐specific critical fire frequency can be predicted, the minimum fire return intervals permitting persistence were 4 yr for U. minor and 3 yr for U. gallii. These patterns are similar to those reported for a range of woody plant species within savanna environments.     

陕北黄土丘陵区撂荒群落共存种活性氧清除指标对水分胁迫的响应

群落共存种对环境变化的生理生态响应差异是群落演替的重要因素,其中活性氧清除是生理生态响应的重要方面。该研究以黄土丘陵区10个常见撂荒群落共存种(阿尔泰狗娃花、猪毛蒿、铁杆蒿、茭蒿、小花棘豆、达乌里胡枝子、白羊草、冰草、中华隐子草和无芒隐子草)为研究对象,测定了不同水分处理条件下(适宜水分、中度胁迫及重度胁迫)生长季中3个月份(8、9和10月)个体和种群的生长表现(个体株高、生物量和种群存活率),及主要活性氧清除指标［类胡萝卜素含量、超氧化物歧化酶(SOD)活性和谷胱甘肽(GSH)含量］,分析群落共存种的生态适合度和活性氧清除指标对水分处理的响应差异及其相互关系,以明确演替生态位置上各指标的变化趋势。结果表明：(1)随水分胁迫程度的增加,演替前期种——猪毛蒿的的适合度有所下降,后期种——白羊草有所上升。(2)GSH和类胡萝卜素含量均随水分胁迫程度增强显著升高。(3)类胡萝卜素含量、SOD活性和GSH含量存在显著的种间差异,其中达乌里胡枝子的类胡萝卜素含量、小花棘豆的SOD活性、中华隐子草的SOD活性和GSH含量、无芒隐子草的类胡萝卜素含量和GSH含量均相对较高,且这3个指标的生长季变化也较为明显,8月份类胡萝卜素含量和GSH含量相对较高,而10月份GSH含量和SOD活性相对较高。(4)演替生态位置上,10个共存种在演替序列上以前期种的适合度和类胡萝卜素含量相对较低,而演替后期种相对较高,表明演替后期的植物较为耐旱。     

Growth characteristics,nutrient allocation and photosynthesis ofCarex species from floating fens

Summary The purpose of this study was to investigate various growth parameters, dry matter and nitrogen, phosphorus and potassium allocation and photosynthesis ofCarex acutiformis, C. rostrata andC. diandra growing in fens with, in this order, decreasing nutrient availability and decreasing aboveground productivity. Plants were grown from cuttings at optimum nutrient conditions in a growth chamber. Growth analysis at sequential harvests revealed that the species had no inherently different relative growth rates which could explain their different productivity, but that their LAR (LWR and SLA) decreased in the orderC. acutiformis, C. rostrata, C. diandra and their NAR increased in this order. All growth parameters decreased during plant growth even under the controlled conditions of the experiment.C. acutiformis allocated relatively much dry matter to the leaves,C. rostrata to the rhizomes andC. diandra to the roots. This may, in part, explain the higher aboveground biomass production ofC. acutiformis in the field. Nitrogen, but not phosphorus and potassium, allocation patterns were different for the three species.C. diandra, the species from the nitrogen-poorest site, had the highest leaf N content of the three species and also a higher chlorophyll content. Related to this, this species had the highest photosynthetic activity of whole plants both when collected from the field and when grown in the growth chamber. The nitrogen productivity was similar for the three species and the photosynthetic nitrogen use efficiency, determined forC. acutiformis andC. diandra, was similar for these two species.C. diandra had the most finely branched root system, i.e., the highest specific root length of the three species and its root surface area to leaf surface area ratio was also the highest. All three species showed higher nitrate reductase activity in the leaves than in the roots when grown on nutrient solution. The growth ofC. diandra at a relatively nutrient-poor site and a rather open low vegetation is assumed to be adapted to its habitat by a relatively high NAR made possible by a high rate of photosynthesis concurrent with a high leaf N content. The growth ofC. acutiformis at a relatively nutrient-rich site and a more dense and higher vegetation is adapted to its habitat by a high LAR.     

The influence of competition from herbaceous vegetation and shade on simulated browsing tolerance of coniferous and deciduous saplings

The ability of saplings to tolerate browsing (i.e. the ability to persist with reduced biomass and to compensate for biomass loss) is influenced by the level of stress and their growth strategies. Ultimately, insight into species‐specific responses of saplings to browsing, shade and competition from neighbours will help explain diversity, structure and function of grazed ecosystems such as the endangered wood‐pasture systems. We measured the survival, whole‐sapling biomass and compensatory growth responses of two coniferous (Picea abies and Abies alba) and two deciduous (Acer pseudoplatanus and Fagus sylvatica) tree species to simulated summer browsing (one single clipping event), shade (installation of a shade cloth) and neighbour removal (mowing surrounding vegetation to ground level) treatments and the interactions between them after two‐growing seasons. For all species, there were interacting effects on growth of browsing and environmental condition (shade and neighbours). Simulated browsing resulted in relatively smaller growth losses when plants were growing slowly due to competitive conditions related to herbaceous neighbours. Although none of the clipped saplings could fully compensate for their biomass losses, the saplings were closer to compensation under high competitive conditions than under low competitive conditions. Survival of the clipped saplings remained relatively high and was only significantly reduced for Picea and Acer. Picea was least tolerant of competition and was the only species for which growth was not negatively affected by strong irradiance of a mountain pasture. Surprisingly, the tolerance of saplings to herbivory as browsing tolerance was enhanced under conditions that negatively affected sapling performance (i.e. survival and growth). Apparently, the relative impact of browsing at the early sapling stage is linked to tree life history characteristics such as competition and shade tolerance and will be lower in situations with intense competitive interactions and/or strong irradiance.