Use of leaf resources by howling monkeys (Alouatta palliata) and leaf-cutting ants (Atta cephalotes) in the tropical rain forest of Los Tuxtlas,Mexico

Use of leaf resources by a troop of howling monkeys and two colonies of leaf cutting ants was studied for an annual cycle in the rain forest of Los Tuxtlas, Mexico. Howling monkeys spent half their annual foraging time feeding on leaves; leaf-cutting ants spent at least 80% of their recorded foraging time harvesting leaves. Both herbivores preferred young leaves over nature ones, and chemical analysis showed that the protein: fibre ratio of the leaves used was correlated with these preferences. Howling monkeys used 34 tree species as leaf sources. Leaf-cutting ants used 40 plant species of which 38 were trees. Eighteen species used by Alouatta were also used by Atta; species of Moraceae and Lauraceae were among the most important in their foraging preferences. The plant species used by monkeys and ants occurred at low densities (? 4.0 ind/ha). The seasonal production of leaves, the high density of leaf-cutting ant colonies at the study site, and the high amounts of young foliage harvested by the ants from tree species, and individual trees used by howling monkeys as sources of young leaves suggest that the foraging activities of Atta may represent a significant pressure upon leaf resources available to Alouatta.     

Comparison of Herbivores and Herbivory in the Canopy and Understory for Two Tropical Tree Species1

The Janzen–Connell model of tropical forest tree diversity predicts that seedlings and young trees growing close to conspecific adults should experience higher levels of damage and mortality from herbivorous insects, with the adult trees acting as either an attractant or source of the herbivores. Previous research in a seasonal forest showed that this pattern of distance‐dependent herbivory occurred in the early wet season during the peak of new leaf production. I hypothesized that distance‐dependent herbivory may occur at this time because the new foliage in the canopy attracts high numbers of herbivores that are limited to feeding on young leaves. As a consequence, seedlings and saplings growing close to these adults are more likely to be discovered and damaged by these herbivores. In the late wet season, when there is little leaf production in the canopy, leaf damage is spread more evenly throughout the forest and distance dependence disappears. I tested three predictions based on this hypothesis: (1) the same species of insect herbivores attack young and adult trees of a given plant species; (2) herbivore densities increase on adult trees during leaf production; and (3) herbivore densities in the understory rise during the course of the wet season. Censuses were conducted on adults and saplings of two tree species, raribea asterolepis and Alseis blackiana. Adults and saplings of both species had largely the same suite of chewing herbivore species. On adults of Q. asterolepis, the density of chewing herbivores increased 6–10 times during leaf production, but there was no increase in herbivore density on adults of A. blackiana. Herbivore densities increased 4.5 times on A. blackiana saplings and 8.9 times on Q. asterolepis saplings during the wet season, but there were no clear trends on the adults of either species. These results suggest that the potential of adult trees as a source of herbivores on saplings depends on the value of new leaves to a tree species' herbivores, which may differ across tree species.     

Seasonal patterns in species richness of herbivores: Macrolepidopteran larvae on Finnish deciduous trees

Abstract. 1. The seasonal distribution of macrolepidopteran species richness on Finnish deciduous trees vaned from positively skewed (peak in spring) to negatively skewed (peak in autumn).
2. The skewness values of species richness had a significant negative correlation ( r = - 0.98) with the duration of the seasonal shoot-growth period of the tree species.
3. Trees which complete their shoot growth early in the season ( Quercus type) produce new leaves only during spring, while trees whose shoot growth continues to autumn ( Populus type) do so throughout the summer.
4. Consequently, there is a difference in the number of available resources in the late summer foliage of different tree species, Trees ceasing leaf production early such as oak ( Quercus robur ) and bird cherry ( Prunus padus ) have one major resource type (mature leaves) in late-season foliage while trees like birches and alders have two Ooung and mature leaves).
5. Because young leaves formed late in the season are preferred to mature ones by some species of herbivores and because other species prefer mature leaves at the same time, the species richness of Populus-type trees is higher later in the season than the species richness of Quercus-type of trees, which have just one type of resource available.     

Interactions among leaf toughness, chemistry, and harvesting by attine ants

Abstract. 1. Young and mature leaves of a tropical legume, Inga edulis var. minutula Schery, are strikingly different in secondary chemistry, especially condensed tannins, and leaf toughness.
2. Bioassays with the two different leaf types indicate that leaf cutter ants, Atta cephalotes (L.), always find mature leaves relatively more acceptable than young leaves when selection was based on chemical cues.
3. Since extracts of young leaves show greziter inhibition of fungal pectinases we suggest that leaf-cutter ants are capable of distinguishing which leaf types are most suitable for the growth of their symbiotic fungus.
4. However, mature leaves are 3 times tougher than young leaves and this prevents leaf-cutter ants from harvesting the more suitable mature leaves.
5. Consequently, bioassays which require cutting before leaf removal indicate that some colonies actually harvest more from the less suitable young leaves.
6. We suggest that the quality of a colony's habitat may indicate whether a colony will harvest more of the less suitable young leaves. Colonies which are harvesting from highly suitable host plants avoid the tropical legume I.edulis while those in poorer habitats accept I.edulis but, because of leaf toughness, mostly harvest the less suitable young leaves.     

Effects of leaf toughness on mandibular wear of the leaf beetle, Plagiodera versicolora

Abstract. 1. The effects of leaf toughness on mandibular wear of the leaf beetle Plagiodera versicolora Laich. (Coleoptera: Chrysomelidae) were evaluated by feeding adult beetles young (tender) and old (tough) leaves of Salix babylonica and S. alba 'Tristis'.
2. Tough leaves erode the cutting surface of beetle jaws more so than tender leaves.
3. Beetles with worn jaws consumed leaves at a slower rate than ones with less mandibular wear.
4. Because rates of leaf consumption and egg production are positively correlated, increased mandibular wear may reduce beetle fecundity.
5. These results support the belief that leaf toughness may act as a potent defence affecting morphology, feeding behaviour, and ultimately spatial and temporal patterns of herbivores.     

Leaf ties as colonization sites for forest arthropods: an experimental study

Abstract.  1. Secondary colonization of leaf shelters constructed by caterpillars has been reported from a number of systems. Both the mechanism (larval or adult movement vs. oviposition) and the cues used by arthropods in locating leaf shelters, however, have received little attention.
2. Artificial leaf shelters (i.e. leaf ties or pairs of leaves clipped together to form sandwiches) were constructed on understorey white oak ( Quercus alba L.) trees and the abundance and species composition of arthropods colonizing and ovipositing on leaf pairs was examined in three treatments: occupied leaf ties (containing a leaf-tying caterpillar), unoccupied leaf ties, and non-tied control leaves.
3. The density of arthropods present in the occupied and unoccupied leaf ties after two weeks was seven and four times greater respectively, than non-tied controls. The guild composition of these early colonists differed among treatments, with the highest densities of leaf chewers, scavengers, and predators in occupied ties and the lowest densities in non-tied controls.
4. The densities of all arthropods ovipositing on leaf pairs in the occupied and unoccupied leaf tie treatments were four and three times greater than non-tied controls. Leaf-chewing insects (including leaf-tiers and non-tying inquiline species) and scavengers showed strong oviposition preferences for tied leaves. However, most species of leaf-tying caterpillars and the psocids (Psocoptera) did not distinguish between occupied and unoccupied ties, suggesting that these groups do not use occupancy-related cues in selecting oviposition sites.     

Growth of an invertebrate shredder on native (Populus) and non-native (Tamarix, Elaeagnus) leaf litter

1. Large-scale invasions of riparian trees can alter the quantity and quality of allochthonous inputs of leaf litter to streams and thus have the potential to alter stream organic matter dynamics. Non-native saltcedar ( Tamarix sp.) and Russian olive ( Elaeagnus angustifolia ) are now among the most common trees in riparian zones in western North America, yet their impacts on energy flow in streams are virtually unknown.
2. We conducted a laboratory feeding experiment to compare the growth of the aquatic crane fly Tipula (Diptera: Tipulidae) on leaf litter from native cottonwood ( Populus ) and non-native Tamarix and Elaeagnus . Tipula showed positive growth on leaf litter of all three species; however, after 7 weeks, larvae fed Tamarix leaves averaged 1.7 and 2.5 times the mass of those fed Elaeagnus and Populus , respectively. Tipula survival was highest on Populus , intermediate on Tamarix and lowest on Elaeagnus .
3. High Tipula growth on Tamarix probably reflects a combination of leaf chemistry and morphology. Conditioned Tamarix leaf litter had intermediate carbon : nitrogen values (33 : 1) compared to Populus (40 : 1) and Elaeagnus (26 : 1), and it had intermediate proportions of structural carbon (42%) compared to Elaeagnus (57%) and Populus (35%). Tamarix leaves are also relatively small and possibly more easily ingested by Tipula than either Elaeagnus or Populus .
4. Field surveys of streams in the western U.S.A. revealed that Tamarix and Elaeagnus leaf packs were rare compared to native Populus , probably due to the elongate shape and small size of the non-native leaves. Thus we conclude that, in general, the impact of non-native riparian invasion on aquatic shredders will depend not only on leaf decomposition rate and palatability but also on rates of leaf litter input to the stream coupled with streambed retention and subsequent availability to consumers.     

Contrasting mechanisms of secondary metabolite accumulation during leaf development in two tropical tree species with different leaf expansion strategies

Young leaves of most species experience remarkably higher herbivore attack rates than mature leaves. Considerable theoretical effort has focused on predicting optimal defense and tradeoffs in defense allocation during leaf expansion. Among others, allocation to secondary chemistry may be dependent on growth constraints. We studied flavanoid production during leaf development in two species of Inga (Fabaceae: Mimosoideae) with different expansion strategies: Inga goldmanii, a species with slowly expanding young leaves, and Inga umbellifera, a species with fast-expanding young leaves. In these two species, the most abundant and toxic class of defensive compounds is flavanoids (which include tannins). We measured their concentration by leaf dry weight, their total content per leaf, their HPLC chemical profile and their toxicity to a generalist herbivore at different expansion levels. Although in both species the flavanoid concentration decreased with increasing leaf expansion, that decrease was twice as pronounced for I. umbellifera as it was for I. goldmanii. I. umbellifera leaves produced flavanoids only during the first half of their development while I. goldmanii leaves continued production throughout. The changes in flavanoid HPLC profiles and toxicity were also more dramatic for I. umbellifera, which had different flavanoids in young than in mature leaves. Relative to I. umbellifera, I. goldmanii showed smaller changes in both flavanoid composition and toxicity in the transition from young to mature leaves. These results indicate that, even though young leaves suffer higher rates of attack and are predicted to have better chemical defenses than mature leaves, growth constraints may modulate defense allocation and thus, evolution of defense strategies.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Seasonal and annual changes in leaf δ13C in two co-occurring Mediterranean oaks: relations to leaf growth and drought progression

1. Changes of δ¹³C and its relation to leaf development, biochemical content and water stress were monitored over a 2 year period in two co-occurring Mediterranean oak species: the deciduous Quercus pubescens and the evergreen Quercus ilex .
2. The time course of leaf δ¹³C showed different patterns in the two species. Young Q. pubescens leaves had a high δ¹³C and a marked decrease occurred during leaf growth. In contrast, leaves at budburst and maturity did not differ significantly in the case of Q. ilex . We suggest that the difference between δ¹³C of young leaves was linked to differential use of reserves of carbon compounds in the two species.
3. δ¹³C values of mature leaves were negatively correlated with minimum seasonal values of predawn water potential, suggesting that a functional adjustment to water resources occurred.
4. There was a significant correlation between individual δ¹³C values for two successive years. This interannual dependence showed that δ¹³C rankings between trees were constant through time.     

Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

Early spring defoliation,secondary leaf flush,and leafminer outbreaks on American holly

Summary Adults of Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, emerge in close synchrony with leaf flush of American holly and feed on and oviposit in soft, partially expanded leaves. Early spring defoliation, such as commonly results from freezing injury to young shoots, is followed several weeks later by a second flush of young leaves from lateral buds. We simulated this phenomenon by manually defoliating whole small trees and individual shoots of large trees to test the hypothesis that freezing injury can encourage leafminer outbreaks by inducing an abundance of soft, protein rich young leaves late in the adult activity period, when availability of vulnerable leaves becomes limited. Defoliation of small trees one or two weeks after bud break resulted in six- to 13-fold increases in the incidence of feeding punctures and larval mines on second flush leaves as compared with densities on original young leaves of control trees. Similarly, we induced significant increases in feeding punctures and larval mines on second flush leaves of individual defoliated shoots, although leaves that did not open until after the flight period escaped this injury. These observations underscore the capability of adult female P. ilicicola to locate and exploit a small number of phenologically available leaves among many hundreds of older leaves on the same tree. By altering the phenology of leaf flush, certain kinds of environmental stress may predispose perennial plants to outbreaks of early season folivores that restrict their feeding or oviposition to very young leaves.     

Abundance and mortality of leaf miners on artificially shaded Emory oak

ABSTRACT.

• 1 We measured solar radiation reaching ten Quercus emoryi Torr. trees and recorded densities of four leaf-mining insect species on these trees from June until September 1982.
• 2 The measurements showed that densities of two leaf miner species were negatively correlated with solar radiation.
• 3 In a field experiment, polypropylene shade fabric was suspended 1.3 m above four experimental trees to test for effects of reduced sunlight.
• 4 Leaves of experimentally-shaded trees were heavier and contained lower percentages of proteins and gallotannins than leaves of control, sun trees, while per cent foliar monomeric, polymeric, and total phenols, and water content did not differ between sun and shaded trees.
• 5 Two of four leaf miner species had greater densities on experimentally-shaded trees than sun trees. One of these two species experienced lower survivorship on sun trees owing to high levels of death from unknown causes on sun trees.
• 6 Two leaf miner species had greater densities on sun trees. One of these species had higher survivorship on shaded trees owing to high levels of parasitism on sun trees.
• 7 We conclude that even subtle differences in shading influences leaf miner density and mortality; however, the effects of shading vary from positive to negative among leaf miner species.

     

Life table analysis of leaf fall in Tasmanian cool temperate forests

The annual leaf fall of three species of cool temperate forest trees was examined for two different consecutive seasons at two localities in Tasmania. The reverse summation of the sequential fall of leaves throughout an annual cycle provided a population biomass that was amenable to life table analyses. Rates of leaf fall were compared between species at different times, and were related to temperature. The threshold temperature for leaf fall to commence was approximately 11–12°C. Estimates of the probability of leaves remaining on trees at different times through the cycle were also obtained.     

The potential for interspecific competition between two abundant defoliators on oak: leaf damage and habitat quality

Abstract. 1. The pedunculate oak, Quercus robur L., suffers high annual levels of spring defoliation in Wytham Woods, Oxon. The two major defoliators, Tortrix viridana L. and Operophtera brumata L., sometimes reach high enough densities to completely defoliate trees.
2. Experiments with larvae of both species enclosed on oak trees in Wytham Woods suggest considerable potential for intra- and interspecific competition between T. viridana and O. brumata.
3. T. viridana and O. brumata consume equivalent amounts of leaf tissue per unit caterpillar density. Despite this, O. brumata is the superior competitor, measured by mortality during the larval feeding period.
4. Evidence is presented that T. viridana larvae depend upon the protected microclimate of leaf rolls to maintain water balance, and that leaf damage may disrupt the construction of, and microclimate within, leaf rolls. O. brumata may therefore be competitively superior because it indirectly interferes with the water balance of T. viridana.
5. The importance of interspecific competition in the dynamics of T. viridana and O. brumata populations is discussed. Stochastic mortality factors acting on first instar larvae of both species probably make intense competition a rare event.     

Seasonality of a tropical leaf-mining moth: leaf availability versus enemy-free space

Abstract.  1. Natural enemies may determine the seasonal occurrence of insect herbivores by restricting the presence of their vulnerable life-stages to periods of low natural enemy activity.
2. A tropical leaf-mining moth, Bucculatrix sp . , feeding on the understorey plant Forsteronia spicata, normally occurs only in the early-wet season in the tropical dry forests of Costa Rica, this period coinciding with a time when natural enemy activity is thought to be lowest and when young leaves are most abundant.
3. A manipulative experiment was run to establish whether the Bucculatrix sp. can undergo another generation in the late-wet season if suitable leaves are present. If the restricted phenology of this species is due to predator avoidance, then its larvae should experience elevated levels of mortality during this late-wet season generation.
4. An artificial second flush of young leaves was induced on F. spicata by stripping them of leaves after the natural first flush of leaves. Bucculatrix sp. larvae were found on the plants induced to flush but none were found on natural plants. No significant difference in pre-adult mortality was found between the larvae on the natural early-wet season and the artificial late-wet season leaves.
5. The primary factor restricting the seasonal occurrence of Bucculatrix sp. is concluded to be the availability of suitable leaves and there is no evidence to suggest that natural enemies play a role in determining its phenology.     

Plant phenology-mediated indirect effects: The gall midge opens the phenological window wider for a leaf beetle

We examined whether larvae of the gall midge Rabdophaga rigidae (Diptera: Cecidomyiidae) can modify the seasonal dynamics of the density of a leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae), by modifying the leaf flushing phenology of its host willow species, Salix serissaefolia and Salix eriocarpa (Salicaceae). To test this, we conducted field observations and a laboratory experiment. The field observations demonstrated that the leaf flushing phenology of the willows and the seasonal dynamics of the beetle density differed between shoots with stem galls and shoots without them. On galled shoots of both willow species, secondary shoot growth and secondary leaf production were promoted; consequently, leaf production showed a bimodal pattern and leaf production periods were 1 to 2 months longer than on non‐galled shoots. The adult beetle density on galled shoots was thus enhanced late in the season, and was found to change seasonally, synchronizing with the production of new leaves on the host willow species. From the results of our laboratory experiment, we attributed this synchrony between adult beetle density and willow leaf flush to beetles’ preference to eat new leaves rather than old. Indeed, beetles consumed five times more of the young leaves when they were fed both young and old leaves. These results indicate that stem galls indirectly enhance the adult beetle density by enhancing food quality and quantity late in the beetle‐feeding season. We therefore conclude that midge galls widen the phenological window for leaf beetles by extending the willows’ leaf flush periods.     

Seasonal and leaf age-dependent changes in occurrence of phyllosphere fungi of giant dogwood

To examine the relative importance of leaf age and season on the occurrence of phyllosphere fungi, temporal patterns of epiphytic and endophytic phyllosphere fungi of giant dogwood (Swida controversa) were studied with reference to leaf emergence at first occurrence and in the middle of the growing season. A total of 15 and 44 species were isolated from the surface and interior of leaves, respectively. On the leaf surface, detection rate of fungi was consistently 100% and their frequencies increased during the growing season, whereas in the leaf interior, detection rate of fungi and their frequencies were low at leaf emergence and gradually increased during the growing season. Six epiphytic and two endophytic fungi were observed frequently. A white sterile mycelium was frequent only on the surface of newly emerged leaves in the first-order shoot in May. The other 7 species increased during the growing season. The frequencies of Phomopsis sp., Pestalotiopsis sp. 1, and Trichoderma viride were higher on the leaves of first-order shoots than those of higher-order shoots that emerged between July and September, suggesting the possible effects of leaf age on their occurrence. On the other hand, the frequencies of Colletotrichum gloeosporioides, Clonostachys rosea, Cladosporium cladosporioides, and Phoma sp. 1 were not different between the first- and higher-order shoots, suggesting the negligible effect of leaf age. The influence of phenological patterns of leaf emergence of deciduous trees on the diversity and composition of assemblages of phyllosphere fungi is discussed.     

Two Mallotus species of different life histories adopt different defense strategies in relation to leaf age

Plants defend their leaves using multiple defense traits that change functions with leaf age. We examined the effects of leaf age on the development of multiple defense traits in two related Mallotus (Euphorbiaceae) species: young plants of the fast‐growing Mallotus japonicus (Spreng.) Müll. Arg. and the slow‐growing Mallotus philippensis (Lam.) Müll. Arg. Sequential leaves of the two species were measured for their leaf area, leaf mass/area, densities of trichomes and pellucid dots, extrafloral nectar volume, and the numbers of extrafloral nectaries and pearl bodies. Mallotus japonicus shifted its defense tactics from direct defense using trichomes and pellucid dots in young leaves to biotic defense using extrafloral nectar and pearl bodies in middle‐aged leaves. In contrast, M. philippensis used direct, chemical defense throughout all leaf ages, together with the shift from indirect, biotic defense using extrafloral nectar in young leaves to direct, physical defense using leaf toughness in middle‐aged leaves. These results strongly suggest that, in relation to life history, plants can alter optimal combinations of multiple defense traits with leaf age.     

Annual photosynthetic activities of temperate evergreen and deciduous broadleaf tree species with simultaneous and successive leaf emergence in response to altitudinal air temperature

To explain why the composition of evergreen and deciduous forests changes along air temperature gradients, we measured several traits of single leaves from temperate deciduous and evergreen broadleaf trees with simultaneous and successive leaf emergence growing at different altitudes in the field. The parameters included seasonal net photosynthetic rate, longevity, mass per area, nitrogen content, and photosynthetic nitrogen-use efficiency. With decreasing altitude, the leaf longevity of deciduous broadleaf trees increased, whereas the maximum net photosynthetic rate decreased. In contrast, leaf longevity of evergreen broadleaf trees decreased, whereas the minimum net photosynthetic rate in winter increased. Along the air temperature gradient, the annual production of deciduous trees with simultaneous leaf emergence may be constant, because the integrated lifetime net photosynthetic rate (ILNPR) of a single leaf changed little. In comparison, deciduous trees with successive leaf emergence may show enhanced annual production with increasing air temperature, by increasing the total leaf number per branch and tree under an extended growing season. Temperate evergreen broadleaf tree species may also show increased annual production with increasing air temperature by sufficiently raising the number of the first-year leaves to the total leaves of branch and tree, which is accelerated by raising the integrated first-year net photosynthetic rate of the single leaf, despite little change in the ILNPR. With increasing air temperature from cool-temperate to warm-temperate zones, evergreen broadleaf tree species gain an advantage of the annual production over deciduous broadleaf tree species with simultaneous leaf emergence.     

Ontogenetic changes in crown architecture and leaf arrangement: effects on light capture efficiency in three tree species differing in leaf longevity

Pronounced strategy shifts along ontogeny have been observed in several tree species, mainly because of the trend to maximize growth during the seedling stage, which constitutes the most vulnerable part of the tree’s life cycle. Our aim here was to analyze the ontogenetic changes in crown characteristics and light capture patterns in three Quercus species: the evergreens Quercus ilex and Quercus suber and the deciduous Quercus faginea co-occurring in a Mediterranean open woodland. The seedlings were distributed in the large clearings among the adults and received full sunlight. We constructed three-dimensional models of the aerial parts of seedlings and mature trees of the three species, using the YplantQMC program. Large differences between growth stages were observed for all variables. The seedlings exhibited smaller branch sizes and crown densities than those observed in the adult trees. Leaf angles to horizontal also tended to increase during ontogeny, whereas leaf dispersion and the observed distances between leaves tended to decrease. The amount of photosynthetic radiation absorbed per unit leaf area throughout the growing season was lower in adult specimens than in young specimens. Changes in absorption efficiency during ontogeny were more intense for the species with longer leaf life span at maturity. We conclude that more intense ontogenetic shifts in species with longer leaf life span reflect the priority change from the maximization of short-term productivity at the seedling stage to maximizing leaf longevity during the adult stage.