The confounding effects of logging on tree seedling growth and herbivory in Central Amazon

Light availability is an important modulator of seedling growth and plant–herbivore dynamics. Logging increases light levels in forests, potentially altering herbivore–plant interactions that drive seedling establishment. We conducted a transplant experiment to evaluate how logging and herbivory affect seedling growth and survival in three shade‐tolerant tree species, at paired canopy gap and understory sites in logged forest and an adjacent unlogged area in central Amazonia (Brazil). Seedlings were either left exposed to naturally occurring insect herbivores or protected from insects by a fine netting structure. We measured the herbivore damage and growth rate of seedlings after 18 mo. In general, logged areas received more light than unlogged sites. Growth and herbivory rates were positively influenced by light, and herbivory was also influenced positively by logging. In gaps, increased growth mitigated foliar damage. Logging resulted in a loss of foliar tissue due to increased herbivory. Herbivory rates were higher in the understory of logged sites than in that of unlogged understory sites, but growth was similar in these areas. Thus, the understory of logged areas provided the least favorable sites for shade‐tolerant tree regeneration, due to higher herbivory rates. The effect of logging on biotic interactions can extend beyond the gaps it creates into untouched understory sites. To our knowledge, this is the first time such a pattern has been observed, highlighting the importance of evaluating the impact of logging on biotic interactions.     

Effects of Light Gaps and Litter Removal on the Seedling Performance of Six African Timber Species1

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split‐plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade‐tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade‐tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade‐tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low‐light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade‐intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade‐intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.     

Insect herbivory declines with forest fragmentation and covaries with plant regeneration mode: evidence from a Mexican tropical rain forest

The consequences of tropical forest fragmentation on herbivory are poorly understood. The limited evidence suggests that forest fragmentation can have positive, negative or neutral effects on herbivory. Inconsistencies may be partly explained by plant interspecific variation and differential responses related to plant life history. In this study we examined the effects of forest fragmentation and plant regeneration mode (shade‐tolerant and light‐demanding species) on sapling herbivory using a large sample of the community (97 species, representing 25% of the woody flora of the study site), and a subset of species shared by forest fragments and continuous forest. For the latter, we also analyzed the effects of species identity on variation in herbivory. Also, for the shared species we used two techniques to measure herbivory: standing herbivory (i.e. instantaneous, actual damage) and cumulative herbivory (i.e. damage, accumulated over time, on initially intact leaves). Insect herbivory was the predominant type of damage in the two forest types, and standing herbivory at both the community and the shared species level was significantly higher in continuous forest than in fragments. Considering shared species, both standing and cumulative herbivory were significantly higher in light‐demanding than in shade‐tolerant species. Cumulative herbivory also showed a significant interaction between forest fragmentation and plant regeneration mode, whereby a significant decline in herbivory in fragments was driven by reduced herbivory in shade‐tolerant species, whereas for light‐demanding species herbivory did not change significantly, due to contrasting species‐specific responses. We conclude that tropical forest fragmentation reduces insect herbivory, but this depends on plant regeneration mode and species identity. These changes could have effects on plant regeneration and diversity in forest fragments via long‐term demographic consequences.     

Gap disturbances in northern old‐growth forests of British Columbia,Canada

Abstract. We characterized the abundance, size and spatial patterning of canopy gaps, as well as gap‐forming processes and light availability in boreal, sub‐boreal, northern temperate and subalpine old‐growth forests of northwestern British Columbia. The proportion of area in canopy gaps ranged from 32% in northern temperate forests to 73% in subalpine forests. Evenly distributed developmental gaps were dominant but permanent openings created by edaphic components and by shrub communities were also common, particularly in subboreal forests. Abundant gaps, large gap sizes, high numbers of gap makers per gap and frequent gap expansion events suggest that gaps have long tenure in these forests. Snapped stems and standing dead mortality were the most common modes of mortality in all forest types resulting in little forest floor disturbance, creating few germination sites for seedling establishment. We found high mean light levels (16–27% full sun) and little difference between non‐gap and gap light environments. Our results suggest that gap dynamics in these forests differ fundamentally from those in temperate and tropical forest ecosystems.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Root/Shoot Allocation and Root Architecture in Seedlings: Variation among Forest Sites, Microhabitats, and Ecological Groups1

I analyzed patterns of variation in root mass allocation and root morphology among seedlings of woody species in relation to environmental factors in four Neotropical forests. Among forests, I explored the response of root traits to sites varying in water or nutrient availability. Within each forest, I explored the plastic response of species to different microhabitats: gaps and understory. Additionally, I explored evidence for life history correlation of root and shoot traits by comparing species differing in their successional group (light‐demanding [22 spp.] or shade tolerant [27 spp.]) and germination type (species with photosynthetic cotyledons or species with reserve cotyledons). At each forest site, young seedlings from 10 to 20 species were excavated. A total of 55 species was collected in understory conditions and 31 of them were also collected in gaps. From each seedling, six morphological ratios were determined. Allocation to roots was higher in forest sites with the lowest soil resources. Roots were finer and longer in the most infertile site, while roots were deeper in the site with the longest dry season. Seedling traits did not differ between germination types. Shade tolerant species allocated more to roots and developed thicker roots than light‐demanding species. Light‐demanding species showed stronger plastic responses to habitat than shade tolerant species, and species with photo‐synthetic cotyledons showed lower plasticity than species with reserve cotyledons. Overall, these results suggest that among Neotropical species, root allocation and root morphology of seedlings reflect plant adjustments to water or nutrient availability at geographic and microhabitat scales. In addition, life history specialization to light environments is suggested by differences among groups of species in their allocation to roots and in their root morphology.     

Primates create seedling growth hotspots through pattern of dung deposition

Primates play important roles in tropical forests through seed dispersal and herbivory. They comprise a large part of the biomass of forest communities and tend to have clumped patterns of defecations (i.e. at favoured food trees or sleeping sites). Therefore, they may also play important roles in accelerating ecosystem nutrient cycling. Here we conduct a controlled growth experiment to quantify the effect of the addition of primate dung on the growth of both light‐demanding and shade‐tolerant seedlings over 1 year in Kibale National Park, Uganda. A mixed model analysis revealed that light‐demanding species were affected by the natural dung treatment and plants with small initial size had accelerated growth, whereas there was no effect on plant growth for shade‐tolerant species. The long‐term implications of increased dung deposition on the local tree community are unclear as shade‐tolerant species may only show an effect over the long‐term and light‐demanding species may only be able to take advantage of the increased growth if subsequently exposed to high light conditions, such as a treefall gap.     

光对热带雨林冠层树种绒毛番龙眼种子萌发及其幼苗早期建立的影响

绒毛番龙眼(Pometia tomentosa)为西双版纳热带季节雨林标志树种,被列为中国珍稀濒危植物。对其开展种子及幼苗生态学研究可为珍稀濒危树种的保护及当地森林恢复提供科学依据。试验采用遮荫试验和野外试验相结合的方法,研究光对绒毛番龙眼种子萌发及幼苗早期建立的影响。结果表明:在遮荫试验30%(林窗中心光)、10%(林窗边缘光)和3.5%(林下光)3个光处理及森林3种生境(林窗中心、林窗边缘和林下)条件下,种子萌发率差异无显著性差异,均达到95%以上,且平均萌发周期小于6 d。3个光处理下的幼苗生长、生物量分配模式及气体交换参数差异显著。30%光下绒毛番龙眼幼苗的根重比(RMR)和茎重比(SMR)最高,10%和3.5%光下幼苗的叶重比(LMR)最高,3.5%光下的叶面积比(LAR)显著高于30%光下。30%光下绒毛番龙眼幼苗的最大净光合速率(P_nmax)、暗呼吸速率(R_d)和光饱和点(I_sat)在3个光处理中均最大,光补偿点(I_comp)则无显著性差异。绒毛番龙眼幼苗具耐荫性,能够在低光环境下长期存活且能缓慢生长;同时具有适应林窗光环境生长的能力,如高的质量相对生长速率(RGR_M)和高度相对生长速率(RGR_H)。林窗的出现是绒毛番龙眼进行成功更新的必要条件,水分可能是限制其幼苗生境选择的另一环境因子。     

Costs versus risks: Architectural changes with changing light quantity and quality in saplings of temperate rainforest trees of different shade tolerance

Light requirements and functional strategies of plants to cope with light heterogeneity in the field have a strong influence on community structure and dynamics. Shade intolerant plants often show a shade avoidance strategy involving a phytochrome‐mediated stem elongation in response to changes in red : far red ratio, while shade‐tolerant plants typically harvest light very efficiently. We measured plant size, stem diameter, internode and leaf lengths in randomly chosen saplings of 11 woody species differing in their shade tolerance in both a secondary forest and an old‐growth temperate evergreen rainforest in southern Chile. We also recorded the irradiance spectrum and the diffuse and direct light availabilities at each sampling point. Significant differences were found for the mean light environment of the saplings of each species, which also differed in basal stem diameter, internode length and leaf length, but not in plant height. Both plant slenderness (plant height/stem diameter) and mean internode length increased with increasing light availability, but no relationship was found between any of these two traits and red : far red ratio. The change in plant slenderness with light availability was of lesser magnitude with increasing shade tolerance of the species, while internode change with light availability increased with increasing shade tolerance of the species. Shade tolerators afford higher costs (thicker stems and plants), which render more biomechanically robust plants, and respond more to the light environment in a trait strongly influencing light interception (internode length) than shade intolerant species. By contrast, less shade‐tolerant plants afforded higher risks with a plastic response to escape from the understorey by making thinner plants that were biomechanically weaker and poorer light interceptors. Thus, species differing in their shade tolerances do differ in their plastic responses to light. Our results contribute to explain plant coexistence in heterogeneous light environments by improving our mechanistic understanding of species responses to light.     

Disturbance regimes,gap‐demanding trees and seed mass related to tree height in warm temperate rain forests worldwide

For tropical lowland rain forests, Denslow (1987) hypothesized that in areas with large‐scale disturbances tree species with a high demand for light make up a larger proportion of the flora; results of tests have been inconsistent. There has been no test for warm temperate rain forests (WTRFs), but they offer a promising testing ground because they differ widely in the extent of disturbance. WTRF is dominated by microphylls sensu Raunkiaer and has a simpler structure and range of physiognomy than tropical or subtropical rain forests. It occurs in six parts of the world: eastern Asia, New Zealand, Chile, South Africa, SE Australia and the Azores. On the Azores it has been mostly destroyed, so we studied instead the subtropical montane rain forest (STMRF) on the Canary Islands which also represents a relict of the kind of WTRF that once stretched across southern Eurasia. We sought to find whether in these six regions the proportion of tree species needing canopy gaps for establishment reflects the frequency and/or extent of canopy disturbance by wind, landslide, volcanic eruptions (lava flow and ash fall), flood or fire. We used standard floras and ecological accounts to draw up lists of core tree species commonly reaching 5 m height. We excluded species which are very rare, very localized in distribution, or confined to special habitats, e.g. coastal forests or rocky sites. We used published accounts and our own experience to classify species into three groups: (1) needing canopy gaps for establishment; (2) needing either light shade throughout or a canopy gap relatively soon (a few months or years) after establishment; and (3) variously more shade‐tolerant. Group 1 species were divided according the kind of canopy opening needed: tree‐fall gap, landslide, lava flow, flood or fire. Only some of the significant differences in proportion of Group 1 species were consistent with differences in the extent of disturbance; even in some of those cases other factors seem likely to have had a major determining influence during evolution. We also sought to determine whether the species that are at least ‘short‐term persistent’ in the soil seed bank (lasting 2–4 years) are all species needing canopy gaps for establishment. The answer was negative; large numbers of seeds of some shade‐tolerants accumulate in the soil, and these species are able to benefit from soil disturbance in deep shade. We found a significant and strong positive relationship in Japan between mean seed mass and mature tree height, a weak positive relationship in New Zealand and no relationship in any of the other four regions. When comparing the seed mass values of Group 1 and Group 3 species we obtained different answers depending on whether or not we confined ourselves to taxonomically controlled contrasts. In only two of the four regions with an appreciable number of species in Group 1 is the mean seed mass of such species significantly lower than that of Group 3 species when taxonomic relatedness is ignored.     

Shade tolerance and herbivory are associated with RGR of tree species via different functional traits

• Relative growth rate (RGR) plays an important role in plant adaptation to the light environment through the growth potential/survival trade‐off. RGR is a complex trait with physiological and biomass allocation components. It has been argued that herbivory may influence the evolution of plant strategies to cope with the light environment, but little is known about the relation between susceptibility to herbivores and growth‐related functional traits.
• Here, we examined in 11 evergreen tree species from a temperate rainforest the association between growth‐related functional traits and (i) species’ shade‐tolerance, and (ii) herbivory rate in the field. We aimed at elucidating the differential linkage of shade and herbivory with RGR via growth‐related functional traits.
• We found that RGR was associated negatively with shade‐tolerance and positively with herbivory rate. However, herbivory rate and shade‐tolerance were not significantly related. RGR was determined mainly by photosynthetic rate (A_max) and specific leaf area (SLA). Results suggest that shade tolerance and herbivore resistance do not covary with the same functional traits. Whereas shade‐tolerance was strongly related to A_max and to a lesser extent to leaf mass ratio (LMR) and dark respiration (R_d), herbivory rate was closely related to allocation traits (SLA and LMR) and slightly associated with protein content.
• The effects of low light on RGR would be mediated by A_max, while the effects of herbivory on RGR would be mediated by SLA. Our findings suggest that shade and herbivores may differentially contribute to shape RGR of tree species through their effects on different resource‐uptake functional traits.

     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Intraspecific variability of specific leaf area fosters the persistence of understorey specialists across a light availability gradient

• Forest understorey plants are sensitive to light availability, and different species groups can respond differently to changing light conditions. A plant trait tightly linked to light capture is specific leaf area (SLA). Studies considering the relative role of within‐ and among‐species SLA variation across different species groups (e.g. specialists and generalists) are rarely implemented in temperate forest understories varying in their maturity.
• We examined community‐level SLA patterns of beech forest understories along a light availability gradient, and for habitat specialists and generalists separately. We then disentangled and quantified the contribution of intraspecific trait variability and interspecific trait differences in shaping SLA patterns.
• We revealed that the increase in community‐level SLA with decreasing light availability was primarily driven by beech forest specialists (and, to a lesser extent, by forest generalists), and this pattern was mainly determined by specialists’ high intraspecific variability. Community‐level SLA was therefore formed by different responses at different organizational levels, i.e. within and among species, and for separate species groups.
• This study provides insights into factors shaping the shade tolerance strategy in beech forest understorey plants; specialists persistence under putative less favourable conditions (i.e. high irradiation) may be fostered by their ability to adjust their light capture strategies intraspecifically.

     

Size‐related life‐history traits in geometrid moths: a comparison of a temperate and a tropical community

1. Comparative studies on insect life histories are facilitated by the increasing availability of reliable phylogenies but are hampered by the scarcity of comparable data. Fortunately, morphological proxies of some life‐history traits can be measured on preserved specimens. 2. This study compared values of size‐related life‐history traits among a tropical (Ugandan) and a temperate (Estonian) assemblage of geometrid moths. 3. A comparative analysis based on an originally derived phylogeny revealed that tropical moths were, on average, larger than temperate ones. Tropical moths also had somewhat lower relative abdomen masses than temperate ones. This indicates that the tropical rather than the temperate moths tend to use an income (rather than capital) breeding strategy. Nevertheless, no difference was found in a related index of pro‐ovigeny. When body size was accounted for, tropical moths were found to lay smaller eggs than temperate ones. 4. The differences between the two compared areas are consistent with selection on higher mobility of the moths imposed by the more diverse tropical vegetation. Relatively larger eggs of temperate moths may constitute an adaptation to overcome the presumably stronger quantitative defences of their host plants. 5. Overall, however, we conclude that the differences in ecologically relevant size‐related traits are relatively low among moth assemblages of a tropical and a temperate forest region, indicating that these environments may not impose radically different selective pressures on insect life histories.     

Plant ecology of tropical and subtropical karst ecosystems

Substantial areas of tropical forests, including those within nine tropical biodiversity hotspots, contain karst landscapes that have developed on soluble carbonate rocks. Here, we review how the ecology of karst forest trees is influenced by hydrological, edaphic, and topographic factors that exhibit fine spatial heterogeneity. Comparative analysis of drought tolerance traits including wood density contributes to the assessment of whether karst tree species are more drought‐tolerant compared to non‐karst trees. Although karst ecosystems are generally considered to have low phosphorus availability, foliar nitrogen‐to‐phosphorus ratios exhibit wide variation across karst regions without a clear difference from non‐karst ecosystems. According to the analyses of leaf phenology, stem water storage, and isotopic signatures from xylem sap, water use strategies of karst trees can be classified into five types: (a) soil water dependent, (b) epikarst water dependent (mainly use water stored in fine pores and gaps within the epikarst rock during the dry season), (c) groundwater dependent, (d) fog water dependent, and (e) drought‐deciduous (shed leaves during the dry season). Overall, published data suggest that only a subset of karst tree species are exclusively distributed within karst hilltops where water availability is limited. The diverse resource acquisition and utilization strategies of karst plants across edaphic habitats must be considered when developing effective strategies to conserve and restore biodiversity in karst landscapes, which are under increasing anthropogenic pressure.     

Why are leaf‐cutting ants more common in early secondary forests than in old‐growth tropical forests? An evaluation of the palatable forage hypothesis

I evaluated the hypothesis that leaf‐cutting ants are more common in early successional forests than in old‐growth forests because pioneer species, which dominate in early successional habitats, appear more susceptible to leafcutters than shade‐tolerant species, which dominate primary forests (palatable forage hypothesis). The relative importance of pioneer and shade‐tolerant species as plant resources for leaf‐cutting ant colonies was evaluated (1) by literature review of leaf‐cutting ants’ diet, and (2) experimentally, using field assays to determine leafcutter's selectivity. Pioneer species were harvested three times more frequently than shade‐tolerant species and made up the largest component of the diet in all the studies reviewed. The amount harvested was not correlated with the plant species abundance. In addition, leaves from pioneer plants were selected eight times more than leaves from shade‐tolerant species in the field assays. These results support the palatable forage hypothesis. Leafcutters probably select pioneer leaves because of their low level of chemical defenses and high nutrient content. The high availability of pioneer species in early successional forest probably decreases the cost to locate palatable resources. Therefore, early successional habitats support more ant colonies than old‐growth forests. On the other hand, the effective defense mechanisms of mature plant species and the high dispersion of palatable plants could explain the low density of leaf‐cutting ant colonies in old‐growth forests. The palatable forage hypothesis is compared with other hypotheses that explain leaf‐cutting ant density. The preference of foundress queens for forest clearings, the dependence of small colonies on herbs, and the importance of pioneer plant species for mature colonies (palatable forage hypothesis) can be considered complementary, because they focus on different stages of the colony's life history. Consequently, the availability of pioneer plants appears to be one of the most influential factors determining mature leaf‐cutting ant nest densities in Neotropical forests.     

Tree dynamics and co‐existence in the montane–sub‐alpine ecotone: the role of different light‐induced strategies

Questions: Is light availability the main factor driving forest dynamics in Pyrenean sub‐alpine forests? Do pines and firs differ in growth, mortality and morphological response to low light availability? Can differences in shade tolerance affect predictions of future biome changes in Pyrenean sub‐alpine forests in the absence of thermal limitation? Location: Montane–sub‐alpine ecotones of the Eastern Pyrenees (NE Spain). Methods: We evaluated morphological plasticity, survival and growth response of saplings of Scots pine, mountain pine and silver fir to light availability in a mixed forest ecotone. For each species, we selected 100 living and 50 dead saplings and measured size, crown morphology and light availability. A wood disk at root collar was then removed for every sapling, and models relating growth and mortality to light were obtained. Results: Fir had the lowest mortality rate (<0.1) for any given light condition. Pines had comparable responses to light availability, although in deep shade Scots pine risked higher mortality (0.35) than mountain pine (0.19). Pines and fir developed opposing strategies to light deprivation: fir employed a conservative strategy based on sacrificing height growth, whereas pines enhanced height growth to escape from shade, but at the expense of higher mortality risk. Scots pine showed higher plasticity than mountain pine for all architectural and morphological traits analysed, having higher adaptive capacity to a changing environment. Conclusions: Our results support the prediction of future biome changes in Pyrenean sub‐alpine forests as silver fir and Scots pine may find appropriate conditions for colonizing mountain pine‐dominated stands due to land‐use change‐related forest densification and climate warming‐related temperature increases, respectively.     

Differential induced response to generalist and specialist herbivores by Lindera benzoin (Lauraceae) in sun and shade

Theoretically, induced defenses should be prevalent within low resource environments like the forest understory where constitutive defenses would be costly. Also, the induced response should be stronger when the herbivore is a generalist rather than a specialist, which often have mechanisms to avoid or overcome plant defenses. These ideas have been previously tested for herbaceous species, and we examined these predictions in Lindera benzoin (spicebush), a common woody shrub of the eastern deciduous forest. Lindera benzoin plants in contrasting light environments served as control plants or were subjected to one of four treatments: application of jasmonic acid, clipping, herbivory by the specialist Epimecis hortaria (tulip tree beauty) and herbivory by the generalist Spodoptera exigua (beet armyworm). Following treatment, we assessed induced responses by measuring leaf chemistry (C/N ratio, protein content, and peroxidase activity), and by using insect bioassays with E. hortaria larvae. We found no difference in peroxidase activity between light environments in controls, plants treated with clipping or jasmonic acid. In plants subject to insect herbivory, peroxidase activity was greater in shade plants than in sun plants. The magnitude of this increase in the shade varied between the herbivores, with a 32 fold increase in plants exposed to the generalist S. exigua and a 9 fold increase in plants exposed to the specialist E. hortaria . Leaves from shade plants had more protein and lower C/N ratios than leaves from sun plants, regardless of induction treatment. In control plants, E. hortaria larvae consumed more leaf biomass and achieved greater final weights in the sun than in the shade, but these differences disappeared with induction treatments were applied. These results are among the first to show rapid induction in a woody plant, and different levels of induction with light environments and with specialist versus generalist herbivores.     

Differences in leaf phenology between juvenile and adult individuals of two tree species in a seasonally dry tropical woodland

In temperate forests, juvenile trees anticipate leaf phenology compared to adults, thus avoiding shading and herbivory. This is also expected to occur in seasonal tropical forests due to intense herbivory and shading during the rainy season; however, the anticipation of leaf phenology by juveniles in seasonal tropical forests has yet to be demonstrated. Stem‐succulent species are expected to be prone to juvenile phenological anticipation because these species are able to use water stored in their stems for leaf flushing in the dry season. We investigated this hypothesis by comparing leaf phenology (bud break, leaf expansion) of juveniles and adults of two species with contrasting wood densities in the transition between dry and rainy seasons in a tropical dry woodland. We also investigated the level of light limitation that juveniles experience in the rainy season. Both species exhibited bud break during the dry season, but only expanded their leaves with the occurrence of the first rains. In general, the stem‐succulent species had a more precocious bud break; however, anticipation by juveniles occurred only in the species with more dense wood. Canopy openness was lower than in temperate deciduous forests, but the fact that the full expansion of leaves occurred only with rainfall indicates that bud break in anticipation of canopy closure contributes only to keeping leaf photosynthetic balance from going negative, and not to higher carbon gain. The importance of anticipated budding for escaping herbivory remains an alternative explanation in need of investigation.