Drivers of invasive tree and shrub natural regeneration in temperate forests

We assessed drivers of ecological success along resource availability gradients for three invasive woody species: Prunus serotina Ehrh., Quercus rubra L. and Robinia pseudoacacia L. We aimed to check how much of invasion success, measured by invader biomass, is explained by propagule pressure and plant community invasibility. Using 3 years of observations from 372 study plots (100 m² each) in temperate forests of Wielkopolski National Park (Poland) we investigated the hierarchy of predictors and partial dependencies using the random forest method. Our study indicated that propagule pressure explained more variance in success of invaders than invasibility—describing availability of resources and competitors in understory vegetation. We also found different responses of seedlings and saplings, connected with dependence on stored carbohydrates, which decreased seedling responses to resource availability gradients. However, resource availability (light and leaf litter predictors) had greater influence than predictors describing understory vegetation. Based on importance and response strength the species studied may be arranged by decreasing requirements for soil fertility and acidity: P. serotina?<?Q. rubra?<?R. pseudoacacia, whereas for light requirements and competition vulnerability the order is: P. serotina?>?Q. rubra?>?R. pseudoacacia. However, low light requirements of R. pseudoacacia may be biased by high proportion of sprouts supplied by parental trees. Results provide guidelines for effective management of invasive woody species in forest ecosystems and describe complex interactions between factors studied on ecological success of invaders.     

<Emphasis Type="Italic">Pinus contorta</Emphasis> invasion into treeless steppe reduces species richness and alters species traits of the local community

Pinus contorta, one of the most invasive tree species in the world, has been proposed as a model species for improving our understanding of invasion ecology. In this study, we assessed the impact of P. contorta invasions on the species richness, diversity and species traits of a resident treeless steppe community. In a Pinus contorta invasion gradient (Patagonia, Chile), we surveyed vegetation from high canopy closure pine invasion to treeless steppe, and computed species richness, diversity and Sørensen similarity indexes. For all species, we determined functional trait values from the literature, data bases, and personal observations. Species richness and diversity were related to canopy cover (a proxy for invasion intensity) using generalized linear mixed-effects models. Changes in species traits due to canopy cover were analyzed using RLQ ordination analysis and the fourth-corner analysis. We found that Pinus contorta canopy cover significantly reduced the number of native species by 70 %, implying a strong effect on species exclusion. A few native species, however, prevail in the novel conditions (e.g. Baccharis magellanica, Acaena integerrima). Species traits changed significantly with increasing pine canopy cover, where P. contorta promoted the existence of traits related to shade-tolerance and conservative reproductive strategies. We conclude that the negative impacts of Pinus contorta into the treeless steppe, including a reduction in the number of species and the shifting to traits adapted to tolerate shade and associated with conservative reproductive strategies, can have severe implications for the conservation of biodiversity and ecosystem functioning where it invades.     

Light energy partitioning,photosynthetic efficiency and biomass allocation in invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> and native <Emphasis Type="Italic">Quercus petraea</Emphasis> in relation to light environment,competition and allelopathy

This study addressed whether competition under different light environments was reflected by changes in leaf absorbed light energy partitioning, photosynthetic efficiency, relative growth rate and biomass allocation in invasive and native competitors. Additionally, a potential allelopathic effect of mulching with invasive Prunus serotina leaves on native Quercus petraea growth and photosynthesis was tested. The effect of light environment on leaf absorbed light energy partitioning and photosynthetic characteristics was more pronounced than the effects of interspecific competition and allelopathy. The quantum yield of PSII of invasive P. serotina increased in the presence of a competitor, indicating a higher plasticity in energy partitioning for the invasive over the native Q. petraea, giving it a competitive advantage. The most striking difference between the two study species was the higher crown-level net CO₂ assimilation rates (A_crown) of P. serotina compared with Q. petraea. At the juvenile life stage, higher relative growth rate and higher biomass allocation to foliage allowed P. serotina to absorb and use light energy for photosynthesis more efficiently than Q. petraea. Species-specific strategies of growth, biomass allocation, light energy partitioning and photosynthetic efficiency varied with the light environment and gave an advantage to the invader over its native competitor in competition for light. However, higher biomass allocation to roots in Q. petraea allows for greater belowground competition for water and nutrients as compared to P. serotina. This niche differentiation may compensate for the lower aboveground competitiveness of the native species and explain its ability to co-occur with the invasive competitor in natural forest settings.     

What makes a plant species specialist in mixed broad-leaved deciduous forests?

Knowledge about relationships between specialization degree of species, i.e. the width of their realized niche and functional traits, may have important implications for the assessment of future population developments under environmental change. In this study, we used a recently introduced method to calculate ecological niche widths of plant species in mixed broad-leaved deciduous forests and to investigate the dependence between niche widths of plants and their functional traits and Ellenberg indicator values. The research is based on a dataset of 4556 phytosociological relevés of mixed broad-leaved deciduous forests in Slovenia. We calculated theta indices for 326 species, which ranks them along a continuous gradient of habitat specialization. For 272 species, we compiled 26 functional traits and Ellenberg indicator values. We found some significant correlations between theta indices of species and their functional traits and Ellenberg indicator values; habitat specialists thrive primarily on the highest altitudes, on colder, dry sites and achieve the age of first flowering later than generalists. They also have smaller seed diameter, lower leaf dry matter content, lower mean canopy height and bigger specific leaf area than generalists. Two species groups, chamaephytes and spring green species, are particularly characterized as specialist species. The added value of our work is in complementing the knowledge about the niche differentiating along different environmental gradients and species coexistence in mixed broad-leaved deciduous forests.     

Interaction between invasive and potentially invasive shrub species does not influence relationships between their ecological success and distance from propagule sources

Although interactions between alien and native plant species are well studied, data on interactions between two co-existing alien species with respect to their invasibility are scarce. The aims of this study were (1) to evaluate three factors shaping abundance of the alien shrub species Cornus alternifolia: abundance of another alien (invasive) shrub species (Prunus serotina), type of tree stand (coniferous vs. broadleaved) and distance to propagule sources and (2) to assess the potential dispersal distance of the species studied. Densities of both species were assessed within 194 experimental plots (located in experimental plantations of trees) in Rogów Arboretum (Central Poland). P. serotina occurred on 79 and C. alternifolia on 33 of the 194 plots. The furthest distance of C. alternifolia from the propagule source was 338 m. C. alternifolia reached higher densities in coniferous than broadleaved tree stands. Density of C. alternifolia depended on tree stand type and distance from the propagule source, but did not depend on density of P. serotina. Density of C. alternifolia decreased with increasing distance from the propagule source; however, this relationship was modified by the type of tree stand: densities were lower in broadleaved than in coniferous stands. The presence of the invasive species seems to neither facilitate nor limit the dispersal distance of C. alternifolia, as these two species differ in shade tolerance. The study also provided the first information about C. alternifolia potential invasiveness, because earlier this species was noticed only as casually escaping from cultivation in Slovakia.     

Intraspecific functional trait variability across different spatial scales: a case study of two dominant trees in Korean pine broadleaved forest

Intraspecific functional trait variability plays an important role in the response of plants to environmental changes. However, it is still unclear how the variability differs across three nested spatial scales (individual, plot, and site) and which determinants (climatic, soil, and ontogenetic variables) shape the trait variability. Along a latitudinal gradient in Korean pine broadleaved forest of northeast China, we quantified the extent of intraspecific variability of four functional traits in two dominant trees Pinus koraiensis and Fraxinus mandshurica at eight sites, including specific leaf area, leaf dry matter content (morphological traits) and leaf nitrogen content, leaf phosphorus content (physiological traits). Results showed a large trait variation within and between species (coefficient variation: 6.07–23.3%). The leaf physiological traits of F. mandshurica and morphological traits of P. koraiensis were more responsive at site scale, while the morphological traits of F. mandshurica and physiological traits of P. koraiensis were more responsive at individual scale. In addition, abiotic and biotic factors explaining functional trait variation differ markedly between the two tree species, with physiological trait of F. mandshurica being more associated with climate and soil, while traits variability in P. koraiensis was not affected by climate, soil, and ontogeny, except for leaf phosphorus content. Overall, we can predict that the physiological traits of broadleaved species tend to be more sensitive to environmental changes, while pines are more sensitive to competition. It is critical to determine which spatial scale and trait type should be taken into account in predictive models of vegetation dynamics.     

Higher phenotypic plasticity does not confer higher salt resistance to <Emphasis Type="Italic">Robinia pseudoacacia</Emphasis> than <Emphasis Type="Italic">Amorpha fruticosa</Emphasis>

A greenhouse experiment was conducted in which two leguminous species commonly used in the Yellow River Delta for vegetation restoration, Robinia pseudoacacia and Amorpha fruticosa, were subjected to five salt treatments: 0, 50, 100, 150, and 200 mmol L^?1. We aimed to determine which of the two species would be better suited for growth in a saline environment, and whether the acclimation capacity to salinity resulted from an inherently higher phenotypic plasticity. The results showed that salinity affected most growth and biomass parameters but had no effects on most leaf traits and physiological parameters of the two species. Height, relative growth rate of crown area, root biomass, and leaf mass ratio of R. pseudoacacia were reduced by higher salinity, while A. fruticosa was not affected. Chlorophyll a-to-chlorophyll b ratio and total antioxidative capacity of A. fruticosa increased with higher salinity, whereas those of R. pseudoacacia remained unchanged. Root mass ratio and vitamin C concentration of both species were not affected by salinity, whereas vitamin C concentration of A. fruticosa was higher than that of R. pseudoacacia. The root-to-shoot ratio of A. fruticosa was higher than that of R. pseudoacacia in most salt treatments. Of all leaf traits, only leaf area differed between treatments. R. pseudoacacia generally exhibited a greater plasticity than A. fruticosa in response to salinity, but A. fruticosa was more resistant to the higher salinities than R. pseudoacacia, and was thus a better candidate for vegetation restoration in saline areas.     

Low impact of disturbance on ecological success of invasive tree and shrub species in temperate forests

Disturbance is claimed to be one of the most important triggers of biological invasions. There is a lack of data about disturbance impacts on the youngest life stage of invasive trees and shrubs. Thus, we aimed to assess the role of disturbances in shaping responses of natural regeneration of three model invasive species—Prunus serotina Ehrh., Quercus rubra L., and Robinia pseudoacacia L.—to disturbances in forest plant communities. Our study was conducted over 3 years on 372 study plots (100 m²), across nine types of temperate forests in Wielkopolski National Park (Poland). Disturbance was assessed using ecological indicator values for disturbance severity and frequency. Our study revealed the high importance of disturbance on species composition of understory vegetation. We also found relationships between ecological success (density and biomass of natural regeneration) of invaders and disturbance indices. These models were statistically significant but their effect sizes were low. Due to the low effect sizes, we can state only limited conclusions about impact of disturbance on ecological success of invasive species natural regeneration. The results suggest that for seedlings (up to 50 cm height—threshold between understory and shrub layer) disturbance, a leading factor in biological invasions of numerous taxa, has a small role in this case. Thus, we may assume that their ecological success is connected with stochastic processes in populations of the invader’s seedlings, rather than with stochastic release from competition caused by disturbances.     

How is light interception efficiency related to shoot structure in tall canopy species?

Coexistence of multiple species is a fundamental aspect of plant and forest ecology. Although spatial arrangement of leaves within crowns is an important determinant of light interception and productivity, shoot structure varies considerably among coexisting canopy species. We investigated the relative importance of structural traits in determining the light availability of leaves (I) and light interception efficiency at the current-year shoot level (LIE_CS; the total light interception of leaves divided by shoot biomass) at the top of crowns of 11 canopy species in a cool-temperate forest in Japan. In accordance with Corner’s rules, the total mass, stem mass, total mass of leaf laminae, individual leaf area, and stem cross-sectional area of current-year shoot were positively correlated with each other, and branching intensity (the number of current-year shoots per branch unit of 1-m length) was inversely correlated with these traits across species. In contrast, I was correlated not with these traits, but with leaf elevation angle (a _L). Moreover, variation in LIE_CS across species was caused by variation in I (thus in a _L). Thus, a _L is a key parameter for the leaf light interception of canopy shoots in this cool-temperate forest. Differences in a _L across species might be related to different physiological strategies that developed in the high light and water-limited environment of forest canopies. Small variation in the length of current-year shoots among species implies that variations in I and LIE_CS would be important for the coexistence of these canopy species.     

Combined effects of climate,habitat, and disturbance on seedling establishment of <Emphasis Type="Italic">Pinus pinaster</Emphasis> and <Emphasis Type="Italic">Eucalyptus globulus</Emphasis>

The natural expansion of forestry trees into habitats outside plantations is a concern for managers and conservationists. We studied seedling emergence and survival of the two main forestry species in Portugal: Eucalyptus globulus (exotic) and Pinus pinaster (native); using a seed addition experiment. Our main objective was to evaluate the combined effects of climate (mild-summer and warm-summer climate), habitat (oak forest and shrubland), and disturbance (vegetation removal and non-disturbance) on the seedling establishment of species in semi- and natural habitats. Furthermore, we tested the effect of the “sowing season” (autumn and spring) on seedling emergence and survival. Overall, seedling establishment of both species was enhanced by light and water. However, we found important interactions among climate, habitat, and disturbance on both species’ emergence and survival. The differences between habitats were more evident in the mild-summer climate than in the warm-summer climate. Our results also suggested that seedling survival may be enhanced by shrub cover in drier conditions (warm-summer climate). Eucalyptus globulus appears more sensitive to drought and disturbance changes than P. pinaster. In shrublands and mild-summer climate conditions, disturbance especially promoted E. globulus seedling establishment, while the forest canopy and the shade appeared to control it in both climatic conditions. After the first summer life, very low seedling survival was observed in both species, although the colonization of new areas appeared to be more limited for E. globulus. Our study suggests that climate conditions influence the effect (direction and intensity) of habitat and disturbance (plant–plant interactions) on seedling survival. Thus, the effect of light availability (forest canopy) and disturbance (vegetation removal) on these species establishment is climate context-dependent. This study presents very useful information to understand future shifts in these species distribution and has direct applications for the management of natural establishment outside the planted areas, and the management of the understorey to favor forest regeneration or limit forest colonization.     

Trait convergence and diversification arising from a complex evolutionary history in Hawaiian species of <Emphasis Type="Italic">Scaevola</Emphasis>

Species variation in functional traits may reflect diversification relating to convergence and/or divergence depending on environmental pressures and phylogenetic history. We tested trait-environment relationships and their basis in finer-scale evolutionary processes among nine extant Hawaiian species of Scaevola L. (Goodeniaceae), a taxon with a complex history of three independent colonizations by different phylogenetic lineages, parallel ecological specialization, and homoploid hybridization events in Hawai‘i. Using a wild population for each species, we evaluated traits related to plant function (morphology, leaf and wood anatomy, nutrient and carbon isotope composition). Hawaiian Scaevola species were distributed across coastal, dry forest and wet forest environments; multivariate environmental analysis using abiotic and biotic factors further showed that species from distantly related lineages inhabited similar environments. Many traits correlated with environment (based on the multivariate environmental analysis), considering both distantly related species and more closely related species. Scaevola species within shared habitats generally showed trait convergence across distantly related lineages, particularly among wet forest species. Furthermore, trait diversification through divergence was extensive among closely related Scaevola species that radiated into novel environments, especially in plant morphology and traits affecting water relations. Homoploid hybrid-origin species were “intermediate” compared to their ancestral parent species, and possessed trait combinations relevant for their current habitat. The diversity in functional traits reflected strong influences of both ecology and evolutionary history in native Hawaiian Scaevola species, and trait correspondence with environment was due to the combination of multiple processes within the taxon: trait pre-adaptation and filtering, evolutionary convergence, divergence, and hybridization.     

Multiple plant traits influence community composition of insect herbivores: a comparison of two understorey shrubs

Structural and nutritional plant traits influence the ability of insect herbivores to locate, consume and persist on their hosts yet it is uncommon for ecologists to consider how multiple plant traits influence insect community composition. We sampled herbivorous insects on two understorey shrub species common to eucalypt forests of south-eastern Australia, namely Cassinia arcuata (Asteraceae) and Daviesia ulicifolia (Fabaceae). Regression analyses were used to assess the relative influence of plant structure (canopy volume), nutritional quality (macronutrients and total phenolics) and plant productivity (leaf litter) on insect abundance and species richness. Total N content of D. ulicifolia was significantly higher than C. arcuata, while the concentrations of P, K, Ca and Mg were higher in C. arcuata. Total phenolics and leaf litter were significantly lower in D. ulicifolia compared to C. arcuata. Insect composition was similar between the two shrubs but C. arcuata supported greater abundances. Canopy volume and the macronutrients P and Ca were important predictors of insect abundance on C. arcuata, whereas canopy volume alone, but neither plant productivity nor macronutrients, influenced the abundance of insects on D. ulicifolia. Ca was an important predictor of insect species richness on C. arcuata and P was an important predictor on D. ulicifolia. By quantifying a range of plant traits, we have provided an understanding of factors likely to influence the composition of herbivorous insects inhabiting these two shrubs. Traits including leaf architecture, foliar morphology and volatile terpenoids may yet explain the greater number of insects on C. arcuata since they influence the availability of microhabitats and apparency of host plants.     

Crown exposure to light and tree allometry of 11 tree species in a snowy cool-temperate forest in Japan

Crown exposure to light (CE) and tree allometry were investigated for 11 species in a snowy cool-temperate secondary forest dominated by Fagus crenata and Betula ermanii in Japan. The 11 species differentiated horizontal and vertical light gradients for regeneration. CE was highly variable across species in small trees, but variation in CE decreased with increasing height. The 11 species were classified into three patterns of height-dependent change in CE in comparison to community-level trends, and rank reversal of CE with increasing height was not apparent. Allometric relationships between trunk diameter (D) and height (H) and between D and trunk length (L) differed little between trees of high and low CE within species. In contrast, slopes of the allometric relationships between D and H differed across species; species with larger maximum height (H _max) were taller at a given D, as was noted in previous studies of warm-temperate and tropical forest trees. Differences in trunk angle among the species of different H _max were the main factor generating the differences in allometric relationships between D and H in this forest. Trunk angle increased with increasing height in the species of large H _max but decreased in those of small H _max. Hence, allometric relationships between D and L were not related to H _max. Since the species of small H _max grow laterally and are easily covered in snow during winter while those of large H _max grow vertically above snow cover, differences in trunk angle may reflect species mechanical properties.     

Refoliation of deciduous canopy trees following severe insect defoliation: comparison of <Emphasis Type="Italic">Fagus crenata</Emphasis> and <Emphasis Type="Italic">Quercus crispula</Emphasis>

Deciduous trees can survive severe defoliation by herbivores and often refoliate in the same season. Refoliation following severe defoliation represents compensatory regrowth to recover foliage biomass. Although the relationship between defoliation intensity and degree of refoliation at the individual level has been quantified following artificial defoliation for saplings and small trees, no study has examined the relationship for canopy trees and interspecific differences in this relationship. In this study, defoliation by gypsy moths in an outbreak year and subsequent refoliation were visually surveyed for canopy trees of Fagus crenata (n?=?80) and Quercus crispula (n?=?113) in central Japan. Defoliation and refoliation estimates were scored in 10% classes as the ratio to foliage present before defoliation. The degree of refoliation and the proportion of refoliated trees were high in severely defoliated trees. For 60 and 100% defoliated trees, respective refoliations were 2 and 66% for F. crenata, and 37 and 88% for Q. crispula. All of the 90 and 100% defoliated trees refoliated. These results indicate that severely defoliated trees show an increased need for refoliation to maintain metabolism. Beta regression analysis showed that Q. crispula possessed higher refoliation capability than F. crenata. This is likely associated with the relatively large storage reserves and recurrent growth flush pattern of oak species, which are strong characteristics of oaks and adaptive for response to herbivory and catastrophic disturbances. Interspecific differences in refoliation capability may exert differential effects on forest ecosystem processes, such as influencing the growth of understory species.     

Stomatal conductance in mature deciduous forest trees exposed to elevated CO<Subscript>2</Subscript>

Stomatal conductance (g _s) of mature trees exposed to elevated CO₂ concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g _s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO₂ compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO₂ treatment. During the 2003 drought in the third treatment year, the CO₂ effect became reversed in Carpinus, resulting in higher g _s in trees exposed to elevated CO₂ compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO₂ enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO₂-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO₂.     

A multi-scale approach to identify invasion drivers and invaders’ future dynamics

Climate, land use and disturbances are well known drivers of invasion. However, their relative influence may change across spatial scales, where climate is expected to be the main filter at broad scales; land use is expected to have more influence at intermediate scales, and disturbance, at fine ones. Understanding the underlying processes that drive invasion patterns at different spatial scales is thus crucial to be able to anticipate the future spread of invaders. Here, we quantified the relative importance of climate, land use, and disturbance on the distribution of the invasive trees Ailanthus altissima and Robinia pseudoacacia, across three nested spatial scales, namely global, country (Spain) and riverbank (three riparian riverbanks). To do so, for each species and scale, we built ensemble species distribution models. We also identified their range filling and inferred the most suitable areas in Spain for them to spread. In general, our study confirms that climate acts as an initial coarse filter of species distribution, whilst both climate and land use were important at the country scale; at the riverbank scale human-mediated disturbances gained importance. However, R. pseudoacacia and A. altissima showed differences in their degree of range filling, where A. altissima has a higher potential for range expansion in the near future. Overall, the integration of different scales into invasion studies shows a great potential to enrich our understanding of species-habitat relationships, and to help anticipate their future dynamics.     

Leaf age dependent changes in within-canopy variation in leaf functional traits: a meta-analysis

Within-canopy variation in leaf structural and photosynthetic characteristics is a major means by which whole canopy photosynthesis is maximized at given total canopy nitrogen. As key acclimatory modifications, leaf nitrogen content (N _A) and photosynthetic capacity (A _A) per unit area increase with increasing light availability in the canopy and these increases are associated with increases in leaf dry mass per unit area (M _A) and/or nitrogen content per dry mass and/or allocation. However, leaf functional characteristics change with increasing leaf age during leaf development and aging, but the importance of these alterations for within-canopy trait gradients is unknown. I conducted a meta-analysis based on 71 canopies that were sampled at different time periods or, in evergreens, included measurements for different-aged leaves to understand how within-canopy variations in leaf traits (trait plasticity) depend on leaf age. The analysis demonstrated that in evergreen woody species, M _A and N _A plasticity decreased with increasing leaf age, but the change in A _A plasticity was less suggesting a certain re-acclimation of A _A to altered light. In deciduous woody species, M _A and N _A gradients in flush-type species increased during leaf development and were almost invariable through the rest of the season, while in continuously leaf-forming species, the trait gradients increased constantly with increasing leaf age. In forbs, N _A plasticity increased, while in grasses, N _A plasticity decreased with increasing leaf age, reflecting life form differences in age-dependent changes in light availability and in nitrogen resorption for growth of generative organs. Although more work is needed to improve the coverage of age-dependent plasticity changes in some plant life forms, I argue that the age-dependent variation in trait plasticity uncovered in this study is large enough to warrant incorporation in simulations of canopy photosynthesis through the growing period.     

Forest Type Influences Population Densities of Nocturnal Lemurs in Manompana,Northeastern Madagascar

Forest loss, fragmentation, and anthropization threaten the survival of forest species all over the world. Shifting agriculture is one of these threatening processes in Madagascar. However, when its cycle is halted and the land is left to regenerate, the resulting growth of secondary forest may provide a viable habitat for folivorous and omnivorous lemur species. We aimed to identify the response of nocturnal lemurs to different successional stages of regenerating secondary, degraded mature, and mature forest across a mosaic-type landscape. We surveyed four nocturnal lemur species (Avahi laniger, Microcebus cf. simmonsi, Allocebus trichotis, and Daubentonia madagascariensis) in four forest types of varying habitat disturbance in northeastern Madagascar. We estimated densities in mature and regenerating secondary forest for the eastern woolly lemur (Avahi laniger) and mouse lemur (Microcebus cf. simmonsi), two sympatric species with folivorous and omnivorous diets respectively. We did not estimate densities of Allocebus trichotis and Daubentonia madagascariensis owing to small sample size; however, we observed both species exclusively in mature forest. We found higher population densities of A. laniger and M. cf. simmonsi in secondary than in mature forest, showing the potential of regenerating secondary forest for lemur conservation. Several environmental factors influenced the detectability of the two lemur species. While observer and habitat type influenced detection of the eastern woolly lemur, canopy height and vine density influenced detection of mouse lemurs. Understanding how different species with different diets interact with anthropogenically impacted habitat will aid future management decisions for the conservation of primate species.     

Response of native Hawaiian woody species to lava-ignited wildfires in tropical forests and shrublands

Wildfires are rare in the disturbance history of Hawaiian forests but may increase in prevalence due to invasive species and global climate change. We documented survival rates and adaptations facilitating persistence of native woody species following 2002–2003 wildfires in Hawaii Volcanoes National Park, Hawaii. Fires occurred during an El Niño drought and were ignited by lava flows. They burned across an environmental gradient occupied by two drier shrub-dominated communities and three mesic/wet Metrosideros forest communities. All the 19 native tree, shrub, and tree fern species demonstrated some capacity of postfire persistence. While greater than 95% of the dominant Metrosideros trees were top-killed, more than half survived fires via basal sprouting. Metrosideros trees with diameters >20 cm sprouted in lower percentages than smaller trees. At least 17 of 29 native woody species colonized the postfire environment via seedling establishment. Although the native biota possess adaptations facilitating persistence following wildfire, the presence of highly competitive invasive plants and ungulates will likely alter postfire succession.     

Spatial and temporal variations in photosynthetic capacity of a temperate deciduous-evergreen forest

Key message

The understory evergreen trees showed maximal photosynthetic capacity in winter, while the overstory deciduous trees showed this capacity in spring. The time lag in productive ecophysiologically active periods between deciduous overstory and evergreen understory trees in a common temperate forest was clearly related to the amount of overstory foliage.

Abstract

In temperate forests, where deciduous canopy trees and evergreen understory trees coexist, understory trees experience great variation in incident radiation corresponding to canopy dynamics represented by leaf-fall and leaf-out. It is generally thought that changes in the light environment affect understory plants’ ecophysiological traits. Thus, to project and estimate annual energy, water, and carbon exchange between forests and the atmosphere, it is necessary to investigate seasonal variation in the ecophysiological activities of both evergreen trees in the understory and deciduous trees that make up the canopy/overstory. We conducted leaf-scale gas-exchange measurements and nitrogen content analyses for six tree species along their heights throughout a complete year. Photosynthetic capacity as represented by the maximum carboxylation rate (V _cmax25) and photosynthetic nitrogen use efficiency (PNUE) of deciduous canopy trees peaked immediately after leaf-out in late May, declined and stabilised during the mid-growing season, and drastically decreased just before leaf-fall. On the other hand, the timing of lowest V _cmax25 and PNUE for evergreen understory trees coincided with that of the highest values for canopy trees. Furthermore, understory trees’ highest values appeared just before canopy tree leaf-out, when incident radiation in the understory was highest. This implies that failing to consider seasonal variation in leaf ecophysiological traits for both canopy and understory trees could lead to serious errors in estimating ecosystem productivity and energy balance for temperate forests.