The influence of large mammalian herbivores on growth form and utilization of mopane trees, Colophospermum mopane, in Botswana's Northern Tuli Game Reserve

Mopane veld is of great value to the general ungulate spectrum in times of drought, and it is capable of retaining its dominance in a community even in the presence of extremely heavy browsing pressure imposed by large browsing mammals. Scrub mopane (hedges) has been regarded as resulting from excessive browsing pressure by large mammals, especially elephants. Both the nutrient and chemical composition of mopane twig bark were investigated, the seasonal results being related back to the seasonal utilization of branches by large mammals. Mopane twigs were most palatable in winter. Eland feed on mopane throughout the year irrespective of palatability. Elephants were rarely present in the scrub mopane area before the onset of spring rains, when the major mopane leaf flush occurs independently of rainfall. The impact of both species was not excessive and recruitment of mopane seedlings does occur. Herbivore browsing is responsible for a mopane morph which buds early and continues to produce accessible, nutritious leaves even when heavily browsed. Many browsing ungulates are reliant on this resource during the stressful transition from spring to summer in south-eastern Botswana.     

Seasonal allocation of defense investment in Ilex opaca Aiton and constraints on a specialist leafminer

Summary Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, is one of the few insect herbivores of American holly. Adult emergence is closely synchronized with leaf flush in spring, and females make numerous feeding punctures on and oviposit in new leaves. Larvae hatch in late May and June, but their feeding period and development are prolonged so that more than 80% of the mine enlargement occurs from January until March of the following year. We propose that this unusual life cycle reflects adaptation to constraints imposed by seasonal and age-related changes in chemical and structural defenses, and in nutritional quality of holly foliage. As holly leaves age, there is a shift in allocation of defense investment away from allelochemicals, including phenolic compounds and saponins, toward leaf sclerophylly, spinose teeth, and low foliar nitrogen and water. Rapid increases in leaf toughness and decreases in nutritional quality limit availability of leaf tissues for adult feeding and oviposition to a two-to threeweek phenological window during leaf flush. Mature holly foliage is a nutritionally poor resource by nearly all criteria known to affect food quality for herbivores. This may be the main reason for the prolonged larval development of P. ilicicola. Alternatively, winter feeding and pupation in spring may be adaptations which help to ensure synchrony of adult emergence with leaf flush. Repeated puncturing by female P. ilicicola does not render leaves more suitable for larvae, nor is it a means by which females sample leaf exudate to assess leaf quality prior to oviposition. Rather, leaf puncturing occurs mostly on leaves that are relatively high in soluble nitrogen, and is apparently a means by which females obtain protein and sugars prior to and during oviposition.The investigation reported in this paper (No. 85-7-8-208) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

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.     

Spring versus autumn leaf colours: Evidence for different selective agents and evolution in various species and floras

Red colouration is common in young and old leaves of broadleaf woody species. Assuming that leaf colours are adaptive, we examined, by comparing the colouration in young versus old leaves, the possibility that different selection agents may have operated on spring versus autumn leaf colouration. We observed spring versus autumn colouration in three very different woody floras (Finland, Japan and Israel) in order to allow for a broad ecological and evolutionary spectrum. The null hypothesis was that if the same selective agents operated in spring and autumn, it is expected that when spring leaves are red, they should always be red in autumn, and when spring leaves are green, they should be green or yellow in autumn. We found that green spring leaves are almost exclusively associated with yellow leaf colour at senescence in autumn. Species with red autumn leaves almost always have at least some red colouration in their spring leaves. However, about half of the species with red spring leaves have yellow autumn leaves. Brown autumn leaves were not common in the species we studied. As about half of the species with red spring leaves have yellow autumn leaves but not vice versa, we conclude that there are many cases in which the selecting agents for spring versus autumn leaf colour were not the same.     

Red young leaves have less mechanical defence than green young leaves

In many plants, leaves that are young and/or old (senescent) are not green. One adaptive hypothesis proposed that leaf color change could be a warning signal reducing insect attack. If leaf coloration involves less herbivory, it remains unclear why leaves in many species are constantly green. To examine whether green leaves reduce herbivory by physical defense as an alternative to the supposed warning signal of red leaves, we conducted comparative analyses of leaf color and protective tissues of 76 woody species in spring. The protective features (trichomes, enhanced cuticle and multiple epidermis) and the distribution of red pigments within leaves were examined in both young and mature leaves. We observed that redness was more frequent in young leaves than in senescent leaves. Compared to 36 species with red young leaves, 40 species with green young leaves showed a significantly higher incidence of enhanced cuticle and trichomes in both phylogenetic and non‐phylogenetic analyses. The phylogenetic analysis indicated that the multiple origins of mechanical protection were generally associated with loss of red coloration. Our finding of relatively poor mechanical protection in red young leaves provides additional evidence for the adaptive explanation of leaf color change.     

Temporal and spatial variations in leaf herbivory within a canopy of<Emphasis Type="Italic"> Fagus crenata</Emphasis>

This study investigated spatio-temporal variation in the leaf area consumed by insect herbivores within a canopy of Fagus crenata, with reference to the light conditions of leaf clusters. There was no clear relationship between photosynthetic photon flux density (PPFD) and consumed leaf area (CLA) in May, immediately after leaf flush, but CLA decreased with an increase in PPFD after June. Leaf mass per area, carbon concentration, C/N ratio, concentration of total phenolics, and condensed tannin concentration were higher in leaves under high light intensity than those of leaves under low light. On the other hand, the nitrogen concentration of leaves decreased as light availability increased. Consequently, within-tree variation in light availability affects the consumption of leaves by insect herbivores through temporal changes in leaf characteristics.     

Developmental Changes in Direct and Indirect Defenses in the Young Leaves of the Neotropical Tree Genus Inga (Fabaceae)

Plant fitness is affected by herbivory, and in moist tropical forests, 70 percent of herbivore damage occurs on young leaves. Thus, to understand the effects of herbivory on tropical plant fitness, it is necessary to understand how tropical young leaves survive the brief, but critical, period of susceptibility. In this study, we surveyed three species of Inga during young leaf expansion. Three classes of toxic secondary metabolites (phenolics, saponins, and tyrosine), extrafloral nectar production, leaf area, and extrafloral nectary area were measured at randomly assigned young leaf sizes. In addition, all defenses were compared for potential trade‐offs during leaf expansion. No trade‐offs among defenses were found, and the concentration of all defenses, except tyrosine, decreased during leaf expansion. We suggest that plants continued to increase phenolic and saponin content, but at a rate that resulted in decreasing concentrations. In contrast, tyrosine content per leaf steadily increased such that a constant concentration was maintained regardless of young leaf size. Nectar production remained constant during leaf expansion, but, because young leaf area increased by tenfold, the investment in extrafloral nectar per leaf area significantly decreased. In addition, nectary area did not change during leaf expansion and therefore the relative size of the nectary significantly decreased during young leaf expansion. These results support the predictions of the optimal defense hypothesis and demonstrate that the youngest leaves have the highest investment in multiple defenses, most likely because they have the highest nitrogen content and are most susceptible to a diversity of herbivores.     

Periods of organogenesis in mono- and bicyclic annual shoots of Juglans regia L. (Juglandaceae)

The organogenetic cycle of shoots on main branches of 4-year-old Juglans regia trees was studied. Mono- and bicyclic floriferous and vegetative annual shoots were analysed. Five parent annual shoot types were sampled between October 1992 and August 1993. Organogenesis of summer growth units was monitored between 16 Jun. and 3 Aug. 1993. Variations over time in the number of nodes, cataphylls and embryonic green leaves of terminal buds were studied. The number of nodes of parent shoot buds was compared with the number of nodes of shoots derived from parent shoot buds. The spring growth units of mono- and bicyclic shoots consist exclusively of preformed leaves which were differentiated, respectively, during the spring flush of growth (mid-April until mid-May) or the summer flush of growth (mid-June until early August) in the previous growing season. Thus, winter buds may consist of flower and leaf primordia differentiated in two different periods during annual shoot extension. The summer growth units of bicyclic shoots consist of preformed leaves that were differentiated in spring buds during the spring flush of growth in the current growing season. Bud morphology is compared between spring and summer shoots.     

Relative growth rate of leaf biomass and leaf nitrogen content in several mediterranean woody species

In many plant species, herbivory is a major determinant of leaf mortality and it can cause a strong reduction in productive potential. Most predation occurs on young, expanding leaves. Thus, a rapid growth of the leaves can reduce the impact of predation. Furthermore, in cold Mediterranean climates the length of the growing season is constrained to a short period in spring and early summer owing both to low winter temperatures and drought stress in early summer. Therefore, a rapid deployment of leaf area and a high photosynthetic capacity during the spring and early summer might have important positive effects on the final carbon balance of the leaf population. Relative growth rates (RGR) of leaf biomass were measured in 19 woody species typical of Central Western Spain with deciduous and evergreen habits. Highly significant differences were detected in the leaf growth rate of the different species. The differences between species, however, did not correlate either with the mean leaf life-span of each of the species or with other leaf traits such as photosynthetic capacity, specific leaf area or nitrogen content. Leaf growth rate was positively correlated with time elapsed between leaf initiation and fruit maturation, so that species with fruit dispersal in spring and early summer in general had lower leaf growth rates than species with autumn fruit shedding. This relationship shows the effects of the concurrence between vegetative and reproductive organs for nutrients and other resources. Nitrogen concentration in the leaves was very high at the time of bud break, and declined during leaf expansion owing to the dilution associated with the increase in structural components. The rate of nitrogen dilution was, thus, positively related to the leaf growth rate. Relative growth rates calculated for nitrogen mass in leaves were very low compared to the growth in total mass. This suggests that most leaf nitrogen is translocated from the plant stores to the leaf biomass before the start of leaf expansion and that the contribution of root uptake during leaf expansion is comparatively low.     

The Herbivore Assemblage, Herbivory and Leaf Chemistry of the Mangrove Kandelia obovata in Two Contrasting Forests in Hong Kong

The herbivore assemblage, intensity of herbivory and factors determining herbivory levels on the mangrove Kandelia obovata (previously K. candel, Rhizophoraceae) were studied over a 13-month period at two forests with contrasting growing conditions in Hong Kong. Mai Po was part of an eutrophic embayment in the Pearl River estuary and generally offered more favourable conditions for mangrove growth, whereas Ting Kok had a rocky substratum and oceanic salinity. Twenty-four insect herbivore species were recorded on K. obovata, with lepidopteran larvae that consume leaf lamina being the dominant species. While leaf litter production was similar at the two forests, herbivory level at Mai Po (mean = 3.9% in terms of leaf area loss) was more severe than that at Ting Kok (mean = 2.3%). Peak herbivory levels were found in summer at both locations (6.5% for Mai Po and 3.8% for Ting Kok). Young leaves of K. obovata at both locations were generally preferred by the herbivores from the period of late spring to summer. Concentrations of most feeding deterrents (ash, crude fibre, and total soluble tannins) were significantly higher in both young and mature leaves at Ting Kok, whereas leaf nutrients (total nitrogen and water) were the same at the two sites. Young leaves at Ting Kok contained about 30% more tannins than their counterparts at Mai Po. Significant differences in leaf chemistry also existed between young and mature leaves at either site. The differences were concomitant with the observed patterns of leaf herbivory on K. obovata, and suggest a potential relationship between environmental quality and plant defence against herbivory.     

Differences in response to simulated herbivory between <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Quercus dentata</Emphasis>

To evaluate the responses of Quercus crispula and Quercus dentata to herbivory, their leaves were subjected to simulated herbivory in early spring and examined for the subsequent changes in leaf traits and attacks by chewing herbivores in mid summer. In Quercus crispula, nitrogen content per area was higher in artificially damaged leaves than in control leaves. This species is assumed to increase the photosynthetic rate per area by increasing nitrogen content per area to compensate leaf area loss. In Quercus dentata, nitrogen content per area did not differ between artificially damaged and control leaves, while nitrogen content per mass was slightly lower in artificially damaged leaves. The difference in their responses can be attributable to the difference in the architecture of their leaves and/or the severeness of herbivory. The development of leaf area from early spring to mid summer was larger in artificially damaged leaves than in control leaves in both species, suggesting the compensatory response to leaf area loss. Leaf dry mass per unit area was also larger in artificially damaged leaves in both species, but the adaptive significance of this change is not clear. In spite of such changes in leaf traits, no difference was detected in the degree of damage by chewing herbivores between artificially damaged and controlled leaves in both species.     

Leaf drop affects herbivory in oaks

Leaf phenology is important to herbivores, but the timing and extent of leaf drop has not played an important role in our understanding of herbivore interactions with deciduous plants. Using phylogenetic general least squares regression, we compared the phenology of leaves of 55 oak species in a common garden with the abundance of leaf miners on those trees. Mine abundance was highest on trees with an intermediate leaf retention index, i.e. trees that lost most, but not all, of their leaves for 2–3 months. The leaves of more evergreen species were more heavily sclerotized, and sclerotized leaves accumulated fewer mines in the summer. Leaves of more deciduous species also accumulated fewer mines in the summer, and this was consistent with the idea that trees reduce overwintering herbivores by shedding leaves. Trees with a later leaf set and slower leaf maturation accumulated fewer herbivores. We propose that both leaf drop and early leaf phenology strongly affect herbivore abundance and select for differences in plant defense. Leaf drop may allow trees to dispose of their herbivores so that the herbivores must recolonize in spring, but trees with the longest leaf retention also have the greatest direct defenses against herbivores.     

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.     

Rhythmic growth and carbon allocation in Quercus robur. Sucrose metabolizing enzymes in leaves

The high sucrose phosphate synthase (SPS) capacity and the low soluble acid invertase activity of mature leaves of the first flush of leaves remained stable during second flush development. Conversely, fluctuations of sucrose synthase (SS) activity were in parallel with the sucrose requirement of the second flush. Sucrose synthase activity (synthesis direction) in first flush leaves could increase in 'response' to sink demand constituted by the second flush growth. Only the ptotosynthates provided by flush mature leaves were translocated for a current flush, while the starch content of these leaves remained stable. After their emergence, second flush leaves showed an increase in SPS and SS (Synthetic direction) activities. The high sucrose synthesis in second flush leaves was used for leaf expansion. When young leaves were 30% fully expanded (stage II20), SPS activity showed little change whereas SS activity declined rapidly toward and after full leaf expansion. The starch accumulation in the young leaves occured simultaneously with their expansion. Developing leaves showed a high level of acid invertase activity until maximum leaf expansion (stage II1). In first and second flush leaves, changes in acid invertase activity correlated positively with changes in reducing sugar concentrations. Alkaline invertase and sucrose synthase (cleavage direction) activities showed similar changes with low values when compared with those of acid invertase activity, especially in second flush leaves. The present results suggest that soluble acid invertase was the primary enzyme responsible for sucrose catabolism in the expanding common oak leaf.     

Do the antiherbivore traits of expanding leaves in the Neotropical tree Inga paraensis (Fabaceae) vary with light availability?

Treefall gaps in tropical forests have a profound effect on plants growing in the understory, primarily due to increased light availability. In higher light, mature leaves typically have increased anti-herbivore defenses. However, since the majority of herbivory occurs while leaves are expanding, it is important to determine whether defense expression during the short period of leaf expansion is canalized (invariant) or plastic in response to variation in light. Therefore, we examined young leaves of Inga paraensis (Fabaceae) saplings growing along a light gradient in a terra-firme forest in Central Amazonia. We quantified leaf production and expansion time, dry mass of phenolics, saponins, and nitrogen, ants attracted to extrafloral nectaries, and leaf consumption. Over the entire light gradient, the number of leaves produced per flush increased by 50?% and the mass of phenolic compounds by 20?%, but no other traits changed. On average, 39?% of leaf area was consumed with no difference across the light gradient. Alone, none of the leaf traits was a significant predictor of leaf consumption, except for phenolics, which showed a positive relationship. Multiple regressions showed that leaf consumption was positively related to more leaves per flush and a higher concentration of phenolics in leaves. Unlike studies of mature leaves, young leaves of I. paraensis show low plasticity in defense traits across a light gradient, suggesting that leaf development is canalized.     

Feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores: The effect of resource abundance and plant chemistry

Summary Leaf tissue preferences of monophagous, oligophagous, and polyphagous insect herbivores were determined using young and mature leaf tissue abundances and herbivore feeding observations. Larvae of monophagous and oligophagous herbivores preferred young leaf tissues while, overall, larvae of polyphagous species preferred mature leaves of their various host plants. Even though a species is often polyphagous over its geographical range, larvae from local populations may be very specialized in their diet. When this occurs these specialized larvae prefer the more nutritious and perhaps more toxic young leaves of some of their host plants. Resource abundance and plant chemistry are discussed as major factors influencing herbivore feeding patterns.     

Leaf Phenology and Leaf Damage of Saplings in the Luquillo Experimental Forest,Puerto Rico1

Changes in light or water availability can result in synchronous leaf production, concentrating food availability for herbivores of young leaves to only a few months. To determine the importance of food availability on herbivory, leaf phenology and leaf damage were studied in the Luquillo Experimental Forest (LEF) of Puerto Rico. We studied 20 individuals of eight species for two years. Every month, new leaves were marked; the following month, leaf area and area of damage were measured. Over two years, comparison of leaf production and percent herbivory were performed for each species, and for all species taken together. More than 30 percent of the annual leaf production occurred in May and June. Leaf production was associated with an increase in PFD (photon flux density) and was not related to the patterns of rainfall. Although leaf production was synchronous, there were no differences in herbivory between the peak and non‐peak periods of leaf production. Possible explanations for the constant levels of herbivory throughout the year are the presence of a generalist herbivore community, the ability of herbivores to track changes in food availability, or high densities of herbivore predators that control herbivore populations.     

Seasonality and edge effect determine herbivory risk according to different plant association models

We report evidence of hierarchical resource selection by large herbivores and plant neighbouring effects in a Mediterranean ecosystem. Plant palatability was assessed according to herbivore foraging decisions. We hypothesize that under natural conditions large herbivores follow a hierarchical foraging pattern, starting at the landscape scale, and then selecting patches and individual plants. A between- and within-patch selection study was carried out in an area formed by scrubland and pasture patches, connected by habitat edges. With regard to between-patch selection, quality-dependent resource selection is reported: herbivores mainly consume pasture in spring and woody plants in winter. Within-patch selection was also observed in scrub habitats, influenced by season, relative patch palatability and edge effect. We defined a Proximity Index (PI) between palatable and unpalatable plants, which allowed verification of neighbouring effects. In spring, when the preferred food resource (i.e. herbs) is abundant, we observed that in habitat edges large herbivores basically select the relatively scarce palatable shrubs, whereas inside scrubland, unpalatable shrub consumption was related to increasing PI. In winter, a very different picture was observed; there was low consumption of palatable species surrounded by unpalatable species in habitat edges, where the latter were more abundant. These outcomes could be explained though different plant associations described in the literature. We conclude that optimal foraging theory provides a conceptual framework behind the observed interactions between plants and large herbivores in Mediterranean ecosystems.     

Chemical and mechanical changes during leaf expansion of four woody species of dry Restinga woodland

Young leaves are preferential targets for herbivores, and plants have developed different strategies to protect them. This study aimed to evaluate different leaf attributes of presumed relevance in protection against herbivory in four woody species (Erythroxylum argentinum, Lithrea brasiliensis, Myrciaria cuspidata, and Myrsine umbellata), growing in a dry restinga woodland in southern Brazil. Evaluation of leaf parameters was made through single-point sampling of leaves (leaf mass per area and leaf contents of nitrogen, carbon, and pigments) at three developmental stages and through time-course sampling of expanding leaves (area and strength). Leaves of M. umbellata showed the highest leaf mass per area (LMA), the largest area, and the longest expansion period. On the other extreme, Myrc. cuspidata had the smallest LMA and leaf size, and the shortest expansion period. Similarly to L. brasiliensis, it displayed red young leaves. None of the species showed delayed-greening, which might be related to the high-irradiance growth conditions. Nitrogen contents reduced with leaf maturity and reached the highest values in the young leaves of E. argentinum and Myrc. cuspidata and the lowest in M. umbellata. Each species seems to present a different set of protective attributes during leaf expansion. Myrciaria cuspidata appears to rely mostly on chemical defences to protect its soft leaves, and anthocyanins might play this role at leaf youth, while M. umbellata seems to invest more on mechanical defences, even at early stages of leaf growth, as well as on a low allocation of nitrogen to the leaves. The other species display intermediate characteristics.     

Landscape nutrition: seeing the forest instead of the trees

Recent theories suggest that herbivores forage across many scales and that foraging decisions are driven by the distribution of nitrogen. However, experimental tests of these predictions across large landscapes are rare and difficult. Pretorius et al. (2011) present an elegant experimental design to test how patch size, local nutrient density and total nutrient load are detected by foraging African elephants (Loxodonta africana) in the Colophospermum mopane shrub veld in South Africa. This experiment should serve as a model for investigations of how herbivores detect and respond to high nutrient patches of different size; it also raises questions for further research, such as the fate of high nutrient patches as elephants disperse nutrients from them in urine, faecal material and carcasses deposited elsewhere in the landscape.