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Summary: Although tannins have been an important focus of studies of plant–animal interactions, traditional tannin analyses cannot differentiate between the diversity of structures present in plants. This has limited our understanding of how different mixtures of these widespread secondary metabolites contribute to variation in biological activity.We used UPLC‐MS/MS to determine the concentration and broad composition of tannins and polyphenols in 628 eucalypt (Eucalyptus, Corymbia and Angophora) samples, and related these to three in vitro functional measures believed to influence herbivore defence: protein precipitation capacity, oxidative activity at high pH and capacity to reduce in vitro nitrogen (N) digestibility.Protein precipitation capacity was most strongly correlated with concentrations of procyanidin subunits in proanthocyanidins (PAs), and late‐eluting ellagitannins. Capacity to reduce in vitro N digestibility was affected most by the subunit composition and mean degree of polymerisation (mDP) of PAs. Finally, concentrations of ellagitannins and prodelphinidin subunits of PAs were the strongest determinants of oxidative activity.The results illustrate why measures of total tannins rarely correlate with animal feeding responses. However, they also confirm that the analytical techniques utilised here could allow researchers to understand how variation in tannins influence the ecology of individuals and populations of herbivores, and, ultimately, other ecosystem processes. [ABSTRACT FROM AUTHOR]

Although ecologists believe that vertebrate herbivores must select a diet that allows them to meet their nutritional requirements, while avoiding intoxication by plant secondary metabolites, this is remarkably difficult to show. A long series of field and laboratory experiments means that we have a good understanding of the factors that affect feeding by leaf-eating marsupials. This knowledge and the natural intraspecific variation in Eucalyptus chemistry allowed us to test the hypothesis that the feeding decisions of greater gliders (Petauroides volans) depend on the concentrations of available nitrogen (incorporating total nitrogen, dry matter digestibility and tannins) and of formylated phloroglucinol compounds (FPCs), potent antifeedants unique to Eucalyptus. We offered captive greater gliders foliage from two species of Eucalyptus, E. viminalis and E. melliodora, which vary naturally in their concentrations of available nitrogen and FPCs. We then measured the amount of foliage eaten by each glider and compared this with our laboratory analyses of foliar total nitrogen, available nitrogen and FPCs for each tree offered. The concentration of FPCs was the main factor that determined how much gliders ate of E. viminalis and E. melliodora, but in gliders fed E. viminalis the concentration of available nitrogen was also a significant influence. In other words, greater gliders ate E. viminalis leaves with a particular combination of FPCs and available nitrogen that maximised the nutritional gain but minimised their ingestion of toxins. In contrast, the concentration of total nitrogen was not correlated with feeding. This study is among the first to empirically show that browsing herbivores select a diet that balances the potential gain (available nutrients) and the potential costs (plant secondary chemicals) of eating leaves. The major implication of the study is that it is essential to identify the limiting nutrients and relevant toxins in a system in order to understand feeding behaviour. [ABSTRACT FROM AUTHOR]

Only about 20 vertebrate species are known to feed regularly on tree sap. One of these is the yellow-bellied glider ( P etaurus australis), a marsupial that obtains between 16% and 80% of its diet from eucalypt sap. We reviewed the literature on sap feeding by the yellow-bellied glider and identified 10 species from which it most frequently obtains sap. These species come mainly from a few sections of the subgenus S ymphyomyrtus, which accounts for two-thirds of over 2000 sap trees reported, but more specifically sections L atoangulatae and M aidenii within this subgenus. Some of these key species contain relatively high concentrations of foliar nutrients and each is an important food tree for at least one of the arboreal folivores - koala ( P hascolarctos cinereus), greater glider ( P etauroides volans), and common ringtail ( P seudocheirus peregrinus) and brushtail possums ( T richosurus vulpecula). Because yellow-bellied gliders, like the arboreal folivores, prefer to feed from certain individuals within a species, we hypothesized that these trees possess a unique chemical signature that links sap-feeding with the concentrations of available nitrogen and formylated phloroglucinol compounds in leaves - the nutritive and defensive chemicals that influence feeding on these species by marsupial folivores. We tested this hypothesis in samples of leaves and bark collected from E . punctata and E . viminalis but found no link between chemistry and sap feeding and conclude that other aspects of an individual tree, such as sap flow or sap chemistry, determine whether gliders will target it for sap. [ABSTRACT FROM AUTHOR]

It is widely postulated that nutritional factors drive bottom-up, resource-based patterns in herbivore ecology and distribution. There is, however, much controversy over the roles of different plant constituents and how these influence individual herbivores and herbivore populations. The density of koala (Phascolarctos cinereus) populations varies widely and many attribute population trends to variation in the nutritional quality of the eucalypt leaves of their diet, but there is little evidence to support this hypothesis. We used a nested design that involved sampling of trees at two spatial scales to investigate how leaf chemistry influences free-living koalas from a low-density population in south east New South Wales, Australia. Using koala faecal pellets as a proxy for koala visitation to trees, we found an interaction between toxins and nutrients in leaves at a small spatial scale, whereby koalas preferred trees with leaves of higher concentrations of available nitrogen but lower concentrations of sideroxylonals (secondary metabolites found exclusively in eucalypts) compared to neighbouring trees of the same species. We argue that taxonomic and phenotypic diversity is likely to be important when foraging in habitats of low nutritional quality in providing diet choice to tradeoff nutrients and toxins and minimise movement costs. Our findings suggest that immediate nutritional concerns are an important priority of folivores in low-quality habitats and imply that nutritional limitations play an important role in constraining folivore populations. We show that, with a careful experimental design, it is possible to make inferences about populations of herbivores that exist at extremely low densities and thus achieve a better understanding about how plant composition influences herbivore ecology and persistence. [ABSTRACT FROM AUTHOR]

The marsupials that eat Eucalyptus in south-eastern Australia provide an example of animals with similar niche requirements occurring sympatrically. They certainly differ in size, ranging from about 1 kg in the greater glider ( Petauroides volans) and the closely related common ringtail possum ( Pseudocheirus peregrinus), to 4 kg (common brushtail possum, Trichosurus vulpecula) and up to 15 kg in the koala ( Phascolarctos cinereus). All species, however, may eat considerable amounts of eucalypt foliage, often favouring the same species, and thus appear to compete for food. In order to better understand the degree of competition for food, we measured feeding by the greater glider in response to increasing concentrations of a specific group of eucalypt plant secondary metabolites (PSM), the sideroxylonals, and then compared it to results published for the other species. The greater glider was more resilient than the other species to increasing concentrations of sideroxylonals. We suggest this allows gliders to feed on leaves from the eucalypt subgenus, Symphyomyrtus, while its small size and gliding ability allow it to feed where koalas cannot, on the young leaves on top of the canopy. In contrast, the common ringtail possum is well adapted to feeding from species of the subgenus Eucalyptus, which do not produce sideroxylonals but contain less available nitrogen (AvailN) than do the symphyomyrtles. These 'nutritional niches' segregate the forest and along with other factors, such as generalist and specialist feeding strategies and differences in body size and requirements for shelter, presumably minimise competition between the marsupial species. [ABSTRACT FROM AUTHOR]

Estimating the nutritional value of a herbivore's diet is difficult because it requires knowing what the animal eats, the relative quality of each component and how these components interact in relation to animal physiology. Current methods are cumbersome and rely on many assumptions that are hard to evaluate. We describe a new method for estimating relative diet quality directly from faeces that avoids the problems inherent in other methods. We combine this method with near infrared reflectance spectroscopy (NIRS) to analyse many samples and thus provide a technique with immense value in ecological studies. The method stems from the correlation between the concentrations of dietary and faecal nitrogen in herbivores eating a tannin-free diet, but a weaker relationship in browsers that ingest substantial amounts of tannins, which form complexes with proteins. These complexes reduce the availability of nitrogen and may increase faecal nitrogen concentrations. Using the tannin-binding compound, polyethylene glycol, we showed that tannin-bound nitrogen is a significant and variable part of faecal nitrogen in wild common brushtail possums ( Trichosurus vulpecula). We developed a technique to measure faecal available nitrogen and found that it predicted the reproductive success of female brushtail possums in northern Australia. Faecal available nitrogen combined with NIRS provides a powerful tool for estimating the relative nutritional value of the diets of browsing herbivores in many ecological systems. It is a better indicator of diet quality than other commonly used single-nutrient measures such as faecal nitrogen and foliage analysis paired with observed feeding behaviour. [ABSTRACT FROM AUTHOR]

Ecologists want to explain why populations of animals are not evenly distributed across landscapes and often turn to nutritional explanations. In seeking to link population attributes with food quality, they often contrast nutritionally positive traits, such as the concentration of nitrogen, against negative ones, such as fibre concentration, by using a ratio of these traits. This specific ratio has attracted attention because it sometimes correlates with the biomass of colobine primates across sites in Asia and Africa. Although empirically successful, we have identified problems with the ratio that may explain why it fails under some conditions to predict colobine biomass. First, available nitrogen, rather than total nitrogen, is nutritionally important, while the presence of tannins is the major factor reducing the availability of nitrogen in browse plant species. Second, tannin complexes inflate measures of fibre. Finally, simple ratios may be unsound statistically because they implicitly assume isometric relationships between variables. We used data on the chemical composition of plants from three continents to examine the relationships between the concentrations of nitrogen, available nitrogen, fibre and tannins in foliage and to evaluate the nitrogen to fibre ratio. Our results suggest that the ratio of the concentration of nitrogen to fibre in leaves does sometimes closely correlate with the concentration of available nitrogen. However, the ratio may give misleading results when leaves contain high concentrations of tannins. The concentration of available nitrogen, which incorporates measures of total nitrogen, dry matter digestibility and tannins, should give a better indication of the nutritional value of leaves for herbivorous mammals that can readily be extrapolated to habitats. [ABSTRACT FROM AUTHOR]

Abstract: Fungi comprise a major part of the diet of many animals. Even so, the nutritional value of fungi has been much debated, with some arguing that fungi are nutritionally poor. However, the chemical composition of fungi and of the biology of the animals that eat them are not well understood, particularly in reference to amino acid (AA) composition of fungi and digestibility of fungal protein. We analysed fibre, total nitrogen (N), available N, and AA contents and measured in vitro digestibility of a wide range of epigeous and hypogeous fungi collected in Australia and the USA to test three hypotheses: (i) fungi are nutritionally poor because they contain few nutrients or are otherwise of low digestibility, (ii) fungi vary substantially in their nutritional composition; and (iii) animals can counter this variable quality by eating diverse taxa. Resultant data indicate many fungi are a reasonable source of AAs and digestible nitrogen. However, they vary highly between species in AA content, and the protein has a poor balance of digestible AAs. This helps explain why many mycophagous animals eat a wide array of fungi and often have digestive strategies to cope with fungi, such as foregut fermentation. Another common strategy is to supplement the diet with high quality protein, such as insect protein. Accordingly, evaluating nutritional value of fungi requires consideration of physiology of the animal species and their whole diet. [Copyright &y& Elsevier]

Abstract: It is becoming increasingly easy to generate genotypic data but much harder to gather an equivalent amount of phenotypic information, particularly for chemical traits. In this study of Eucalyptus globulus ssp. globulus, we measured about 60 chemical leaf traits of trees growing in a common garden to address the following questions: (1) how much variation is there between geographic regions, populations within regions and within populations? (2) How do chemical traits vary over the species’ geographic range? (3) If so, does E. globulus ssp. globulus exhibit distinct chemotypes – plants that are morphologically similar but which differ chemically? (4) Are the affinities between E. globulus ssp. globulus and closely related subspecies apparent in the chemical variation? Variation among trees within populations contributed most variation in leaf chemistry followed by variation between geographic regions. For many traits, variation among populations within proposed races and variation among proposed races within geographic regions explained little of the total variation. There was a cline in the concentration of secondary chemicals with the lowest concentrations in Tasmanian populations and the highest in those from eastern Victoria, with intermediate concentrations in populations from Bass Strait Islands. We identified three chemotypes, characterised by specific terpenes and formylated phloroglucinol compounds. The frequency of occurrence of these chemotypes showed a geographic pattern also, with “chemotype 1” predominating in Tasmania, while “chemotypes 2 and 3” occurred at highest frequency in eastern Victoria. We suggest that the chemotypes reflect introgression between E. globulus ssp. globulus and the three closely related subspecies –E. globulus ssp. bicostata, E. globulus ssp. maidenii and E. globulus ssp. pseudoglobulus. Although the formation of land-bridges with fluctuating sea levels has no doubt shaped the evolutionary history of all four subspecies, we propose that the migratory swift parrot (Lathamus discolor), an important pollinator and a species closely associated with E. globulus, has helped shape the evolution of the four tree subspecies. [Copyright &y& Elsevier]

Summary [ABSTRACT FROM AUTHOR]

The chemical quality of forage may determine landscape use and habitat quality for some herbivorous species. However, studies that investigate the relationship between foliar chemistry and foraging choices in wild vertebrates are rare. Petauroides volans (the greater glider) is unique among Australian marsupial folivores because it glides. It also frequently consumes foliage from both major Eucalyptus subgenera, Eucalyptus (common name 'monocalypt') and Symphyomyrtus (common name 'symphyomyrtle'), which differ markedly in their foliar chemistry. Such differences are thought to be a product of co-evolution that also led to guild-specific plant secondary metabolite (PSM) specialization among other marsupial eucalypt folivores. To explore whether foliar chemistry influences tree use, we analyzed foliage from eucalypt trees in which we observed P. volans during a radio tracking study and from eucalypt trees in which animals were never observed. We used a combination of chemical assays and near infrared spectrophotometry (NIRS) to determine concentrations of nitrogen (N), in vitro available nitrogen (AvailN), and in vitro digestible dry matter (DDM) from foliage sampled from the monocalypt and symphyomyrtle species, and total formylated phloroglucinol compounds (FPCs) and sideroxylonals (a class of FPCs) from the symphyomyrtle species (FPCs do not occur in monocalypts). Tree size and spatially-dependent, intraspecific variations in sideroxylonals and DDM concentrations in the symphyomyrtle foliage and of N, AvailN, and DDM in the monocalypt species were important indicators of tree use and habitat suitability for P. volans. The results i) demonstrate that guild-specific PSMs do not always lead to guild-specific foraging; ii) provide a compelling co-evolutionary case for the development of gliding in P. volans; and iii) have implications for the management and conservation of this and other folivorous species. [ABSTRACT FROM AUTHOR]

Different folivorous marsupials select their food from different subgenera of Eucalyptus, but the choices cannot be explained by known antifeedants, such as formylated phloroglucinol compounds or tannins, or by nutritional quality. Eucalypts contain a wide variety of plant secondary metabolites so it is difficult to use traditional methods to identify the chemicals that determine food selection. Therefore, we used a metabolomic approach in which we employed 1H nuclear magnetic resonance spectroscopy to compare chemical structures of representatives from the two subgenera and to identify chemicals that consistently differ between them. We found that dichloromethane extracts of leaves from most species in the subgenus Eucalyptus differ from those in Symphyomyrtus by the presence of free flavanones, having no substitution in Ring B. Although flavanoids are known to deter feeding by certain insects, their effects on marsupials have not been established and must be tested with controlled feeding studies. [ABSTRACT FROM AUTHOR]

Background and Aims Both environmental and genetic effects contribute to phenotypic variation within and among populations. Genetic differentiation of quantitative traits among populations has been shown in many species, yet it can also be accompanied by other genetic changes, such as divergence in phenotypic plasticity and in genetic variance. Sideroxylonal (a formylated phloroglucinol compound or FPC) is an important chemical defence in eucalypts. The effect of environmental variation on its production is a critical gap in our understanding of its genetics and evolution. Methods The stability of genetic variation in sideroxylonal was assessed within and among populations of Eucalyptus tricarpa in three replicated provenance/progeny trials. The covariance structure of the data was also modelled to test whether genetic variances were consistent among populations and Fain’s test was applied for major gene effects. Key Results A significant genotype × environment interaction occurred at the level of population, and was related to temperature range and seasonality in source populations. Within-population genetic variation was not affected by genotype × environment effects or different sampling years. However, within-population genetic variance for sideroxylonal concentration differed significantly among source populations. Regression of family variance on family mean suggested that this trait is subject to major gene effects, which could explain the observed differences in genetic variances among populations. Conclusions These results highlight the importance of replicated common-garden experiments for understanding the genetic basis of population differences. Genotype × environment interactions are unlikely to impede evolution or responses to artificial selection on sideroxylonal, but the lack of genetic variation in some populations may be a constraint. The results are broadly consistent with localized selection on foliar defence and illustrate that differentiation in population means, whether due to selection or to drift, can be accompanied by changes in other characteristics, such as plasticity and genetic variance. [ABSTRACT FROM AUTHOR]

Abstract: This paper provides a synthesis of the evidence on the patronage growth performance of bus improvement measures in urban settings. The evidence includes a summary of experience in Europe, North America and Australasia focusing on service improvement measures including network structure and service levels, bus priority measures, vehicles and stop infrastructure, fares and ticketing systems, passenger information and marketing, personal safety and security and synergy effects of measures. The source is the research literature and documented experienced from a series of studies undertaken by the authors over the last decade. It includes the results of an international bus expert ‘Delphi’ survey concerning bus improvement measures focussed on patronage growth. The paper synthesises the evidence to identify measures which are most likely to grow patronage including consideration of cost-effectiveness of measures. [Copyright &y& Elsevier]

Many regard the concentrations of nitrogen (N), tannins and plant cell wall constituents (fibre) as key indicators of food quality and habitat suitability for browsing herbivores; yet there is no method for measuring their combined effects. We have developed a simple in vitro assay for measuring the effects of tannins and fibre on N availability in browse. We determined the effects of tannins by measuring the polyethylene glycol (PEG)-binding capacity (PEG-BC) of Eucalyptus leaf samples, followed by a two-stage in vitro digestion with pepsin and cellulase to determine the digestibility of dry matter and N. There was a significant relationship between concentrations of digestible N and the PEG-BC of the leaves. Furthermore, adding PEG significantly improved the digestibility of N. Our results concur with in vivo observations from several mammalian species. This suggests that our method is effective for measuring the nutritional quality of browse and the benefits of adding PEG, providing some index of the detrimental effects of tannins. We further simplified the assay by removing the PEG step, allowing us to quickly analyse samples in bulk. Nevertheless, this simplified method is still not practical for analysing the many samples necessary to compare the nutritional values of different tracts of forest. We used near-infrared reflectance spectroscopy to produce calibration equations and predicted total and digestible N in 322 trees at eleven sites. Both within and between sites, we found a wide variation in concentrations of digestible N but a much lower variation in total N, with either no relationship or poor relationships between the two measures. This confirms the variability in the nutritional quality of eucalypt forests, which may explain the distribution and abundance of mammalian herbivores. Thus, our assay provides a useful tool for understanding how food resources influence herbivore populations at different scales. [ABSTRACT FROM AUTHOR]

In a given period of time, herbivores often eat less as dietary plant secondary metabolite (PSM) concentrations increase. This reduction in total food intake is interpreted as a need for the herbivore to regulate PSM ingestion in order to avoid toxication. However, regulation of PSM ingestion involves more than the reduction of total intake; it involves an alteration of meal patterns, through a reduction in the number and/or the size of the meals eaten. Despite this, studies of how herbivores alter their meal patterns when offered varying concentrations of PSMs are rare. We investigated whether koalas adjust the number and/or the size of their meals when offered eucalypt foliage varying naturally in concentrations of formylated phloroglucinol compounds (FPCs), a group of PSMs that have previously been shown to inhibit total food intake. High FPC concentrations caused koalas to eat more slowly, eat shorter meals and eat less per meal, which resulted in a reduced total intake. Because increasing FPC concentrations did not cause koalas to alter the number of meals that they ate, clear individual differences between koalas were observed, where some consistently ate fewer larger meals and others ate many smaller meals. Thus, different feeding strategies may still achieve the same outcome of a regulated intake of PSMs. The changes observed match the meal patterns of other herbivores ingesting PSMs known to stimulate nausea and emetic pathways, supporting the idea that feedback signals from nausea are an important way that koalas avoid toxication when eating eucalypt foliage. [ABSTRACT FROM AUTHOR]

Plants encounter a broad range of natural enemies and defend themselves in diverse ways. The cost of defense can be reduced if a plant secondary metabolite confers resistance to multiple herbivores. However, there are few examples of positively correlated defenses in plants against herbivores of different types. We present evidence that a genetically variable chemical trait that acts as a strong antifeedant to mammalian herbivores of Eucalyptus also deters insect herbivores, suggesting a possible mechanism for cross-resistance. We provide field confirmation that sideroxylonal, an important antifeedant for mammalian herbivores, also determines patterns of damage by Christmas beetles, a specialist insect herbivore of Eucalyptus. In a genetic progeny trial of Eucalyptus tricarpa, we found significant heritabilities of sideroxylonal concentration (0.60), overall insect damage (0.34), and growth traits (0.30–0.53). Population of origin also had a strong effect on each trait. Negative phenotypic correlations were observed between sideroxylonal and damage, and between damage and growth. No relationship was observed between sideroxylonal concentration and any growth trait. Our results suggest that potential for evolution by natural selection of sideroxylonal concentrations is not strongly constrained by growth costs and that both growth and defense traits can be successfully incorporated into breeding programs for plantation trees. [ABSTRACT FROM AUTHOR]

Most herbivores eat more and survive better when they have access to a variety of foods. One explanation involves the detoxification of plant secondary metabolites (PSMs). By feeding from a variety of plants that contain different classes of PSMs, animals can use multiple detoxification pathways and presumably consume more food. Although popular, this theory is difficult to test because it requires knowledge of the detoxification pathways of each PSM in the diet. We established that common brushtail possums (Trithosurus vulpecula) use various combinations of oxidation, hydrolysis, and conjugation with glucuronic acid (GA) or glycine to detoxify six PSMs. Compared to their ingestion of a single PSM, possums ate more when offered a choice between two diets containing PSMs that require apparently independent detoxification pathways (benzoate and 1,8-cineole, benzoate and p-cymene, benzoate and orcinol, benzoate and salicin, or orcinol and 1,8-cineole). However, possums still did not eat as much of these diets as they did of a basal diet free of PSMs. This suggests that detoxification pathways are never independent, but are separated instead by degrees. In contrast, possums offered a choice of two PSMs that require competing detoxification pathways (1,8-cineole and p-cymene, 1,8-cineole and salicin, or orcinol and salicin) ate no more than when offered diets containing one of the compounds. There was an exception: even though both rutin and orcinol are detoxified via conjugation with GA, the feeding behavior of possums did not suggest competition for detoxification pathways. This implies that the supply of GA is not limiting. This study provides the first convincing evidence that herbivorous mammals can eat more by selecting mixed diets with a diversity of PSMs that make full use of their detoxification potential. It also emphasizes that other behavioral and physiological factors, such as transient food aversions, influence feeding behavior. [ABSTRACT FROM AUTHOR]

A central theory in mammalian herbivore-plant interactions is that feeding decisions depend on an animal's capacity to detoxify plant secondary metabolites (PSMs). Thus, if a herbivore could detoxify a PSM faster, it might eat more. This theory has been difficult to test because the mechanisms by which herbivores detoxify most PSMs are largely unknown. We investigated whether common brushtail possums alter aspects of their feeding in response to an increased capacity to detoxify benzoic acid, a common PSM. Animals detoxify benzoic acid primarily by conjugating it with glycine to form benzoyl glycine (hippuric acid). Therefore, adding glycine to a diet that contains benzoic acid provides a simple way to study how detoxification constrains feeding in mammals. Brushtail possums offered supplementary glycine metabolized benzoate faster and, in response, ate more. Furthermore, when given a choice, brushtail possums selected a diet containing both benzoate and glycine over diets with a high concentration of just one of these supplements. The possums' aversion to glycine may explain why their ability to select a favorable diet combination from feeders containing diets with each of the supplements seemed less acute: they preferred diets containing benzoate alone over those containing glycine and did not mix their diet even though it would have allowed them to eat more benzoate and thus more food. Administering the anti-emetic drug, ondansetron (a specific 5-HT3 receptor antagonist), did not alter the amount eaten of diets containing benzoate, glycine, or both, suggesting that 5-HT3 receptors do not regulate benzoate or glycine intake. The results show that common brushtail possums recognize changes in their detoxification capacity and alter their feeding response accordingly. [ABSTRACT FROM AUTHOR]

The distribution of marsupial folivores in Australian eucalypt forests is extremely patchy and, although distribution patterns are known to be partly determined by nutritional factors, the role of plant secondary metabolites (PSMs) has been little studied. We investigated variation in the foliar concentrations of five constituents: total phenolics (TP); the foliar component bound by polyethylene glycol (PEGBC); the formylphloroglucinol compound, sideroxylonal (a strong deterrent of mammalian herbivory); the terpene, cineole; and nitrogen (N). These constituents were analyzed using near infrared spectroscopy in 501 individual tallowwood (Eucalyptus microcorys) trees at 42 sites across a broad geographic range in eastern Australia. We tested three hypotheses: (1) trees with lower foliar N produce higher PSM concentrations; (2) PSM concentrations are greater at colder sites; and (3) PSM concentrations are higher at sites with low resource availability. Hypothesis 1 was true for all PSMs considered and was the result of variation that occurred within sites. This is consistent with the carbon/nutrient balance hypothesis and with the hypothesis that PSMs protect leaves from photodamage. Hypothesis 2 was upheld for all constituents. This was consistent with both the growth-differentiation balance hypothesis, and with the hypothesis that PSMs represent an evolutionary adaptation to greater risks of photodamage and frost damage in cold environments. Hypothesis 3 was rejected, with one PSM, sideroxylonal, being affected by resource availability in a manner opposite to that predicted. The optimal defense hypothesis offers an explanation for this result (as sideroxylonal concentrations increased along with the apparent nutritional quality of the foliage, possibly to provide defense against higher rates of herbivory). We found that trees associated with koala fecal pellets had lower concentrations of sideroxylonal and cineole. We discuss this finding and further implications for the herbivores of Eucalyptus of the patterns of plant defense that we detected. [ABSTRACT FROM AUTHOR]

Explores the problems surrounding intercessory prayer and the centrality of Christ's mediatory function. Attempts to re-appropriate the language of intercession within a spirituality; Relationship between Jesus and intercession; Theological significance of much of Jesus' ministry and death; Jesus' paradigmatic prayer of intercession in Gethsemane; Christologizing of the Spirit.

Ecologists have long been interested in how the nutritional composition of leaves changes as they age, and whether this affects herbivore feeding preferences. As a consequence, the literature abounds with reports that younger leaves contain higher concentrations of nitrogen and plant secondary metabolites (PSMs) than do older leaves. Most of these studies, however, base their conclusions on average values that often mean little to herbivores. We examined this issue in the well-studied marsupial-eucalypt system, using Eucalyptus melliodora and captive common brushtail possums (Trichosurus vulpecula) offered branches from individual trees containing both young and mature leaves. Like many plants, the concentrations of N and PSMs differed among individual E. melliodora. Although young leaves were, on average, “better defended” by the PSM sideroxylonal than were mature leaves, some trees produced leaves that were relatively undefended at both ages. In response, possums chose different proportions of young and mature leaves depending on the chemistry of the individual tree. Possums did not always prefer leaves with lower concentrations of sideroxylonal (mature leaves) or those with higher concentrations of available N (young leaves). Instead, the sideroxylonal concentration of young leaves dictated their choice: possums preferred young leaves with low sideroxylonal concentrations, but not with high concentrations. By skewing their feeding toward trees producing young leaves with low concentrations of PSMs, possums may influence plant fitness. Researchers will detect these potentially important interactions only if they are aware that measuring variation among plants discloses more information than do average relationships. [ABSTRACT FROM AUTHOR]

Adequate nutrition is a fundamental requirement for the maintenance and growth of populations, but complex interactions between nutrients and plant toxins make it difficult to link variation in plant quality to the ecology of wild herbivores. We asked whether a 'foodscape' model of habitat that uses near-infrared spectroscopy to describe the palatability of individual trees in the landscape, predicted the foraging decisions of a mammalian browser, the koala ( Phascolarctos cinereus). Specifically, we considered four behavioural decision points at which nutritional quality may influence an animal's decision. These were: which tree to enter, whether to feed from that tree, when to stop eating, and how long to remain in that tree. There were trends for koalas to feed in eucalypt trees that were more palatable than unvisited neighbouring conspecific trees, and than trees that they visited but did not eat. Koalas ate longer meals in more palatable trees, and stayed longer and spent more time feeding per visit to these trees. Using more traditional chemical analyses, we identified that an interaction between the concentrations of formylated phloroglucinol compounds (a group of plant secondary metabolites) and available N (an integrated measure of tannins, digestibility and N) influenced feeding. The study shows that foodscape models that combine spatial information with integrated measures of food quality are a powerful tool to predict the feeding behaviour of herbivores in a landscape. [ABSTRACT FROM AUTHOR]

Mantled howler monkeys ( Alouatta palliata) occupy a wide variety of tropical habitats and are the most folivorous of New World primates. However, their diet may include fruits, buds, petioles, and flowers, as well as leaves, suggesting they must cope with variations in the nutrient composition of their food. We studied the physiological basis of the dietary flexibility of these monkeys by comparing food choice, digestive performance and patterns of digesta flow in six adults, fed diets of either leaves or a mixture of fruit and leaves. Although monkeys ate similar amounts of the two diets, they ingested more digestible protein when offered the leaf diet, on which they lost body mass, but they ingested much more soluble sugars when offered fruit and leaves on which they gained mass. Digestibilities of dry matter, fat, energy and fibre did not differ between diets, but those of crude protein, soluble sugars and minerals were higher on the fruit–leaf diet. Mean retention times in the gut of solute (Co-EDTA) and particulate markers (Cr-mordanted cell walls) did not differ between diets, but on both diets the monkeys retained the particulate marker (mean retention time ca 55 h) for longer than they did the solute marker (MRT ca 50 h). A lack of selective retention of solutes and small particles in the gastro-intestinal tract of howler monkeys probably restricts them to mixed diets but their digestive strategy is sufficiently flexible to allow them to feed on a diet of leaves when fruit is unavailable. [ABSTRACT FROM AUTHOR]

The detoxification limitation hypothesis predicts that the metabolism and biotransformation of plant secondary metabolites (PSMs) elicit a cost to herbivores. There have been many attempts to estimate these costs to mammalian herbivores in terms of energy, but this ignores what may be a more important cost-increases in protein turnover and concomitant losses of amino acids. We measured the effect of varying dietary protein concentrations on the ingestion of two PSMs (1,8 cineole-a monoterpene, and benzoic acid-an aromatic carboxylic acid) by common brushtail possums ( Trichosurus vulpecula). The dietary protein concentration had a small effect on how much cineole possums ingested. In contrast, protein had a large effect on how much benzoate they ingested, especially at high dietary concentrations of benzoate. This prompted us to measure the effects of dietary protein and benzoate on whole-body protein turnover using the end-product method following an oral dose of [N] glycine. Increasing the concentration of dietary protein in diets without PSMs improved N balance but did not influence whole-body protein turnover. In contrast, feeding benzoate in a low-protein diet pushed animals into negative N balance. The concomitant increases in the rates of whole-body protein turnover in possums eating diets with more benzoate were indicative of a protein cost of detoxification. This was about 30 % of the dietary N intake and highlights the significant effects that PSMs can have on nutrient metabolism and retention. [ABSTRACT FROM AUTHOR]

Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes., We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences., Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant ( P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis ( PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes., Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species. [ABSTRACT FROM AUTHOR]

Mosaic trees contain more than one phenotype. The two Eucalyptus mosaic trees studied here ( E. melliodora and E. sideroxylon) are predominantly susceptible to insect herbivory, with the leaves on a single large branch on each tree resisting herbivory. We used gas chromatography-mass spectrometry and high-pressure liquid chromatography to analyze the chemical profile of leaves of the mosaic trees, as well as leaves of adjacent non-mosaic con-specifics. We show that the leaves of the two phenotypes are distinctly different. Compared to the susceptible (S) leaves on the same tree, the resistant (R) leaves on the mosaic trees had low concentrations of sesquiterpenes ( E. melliodora: 2 vs. 24 mg·g dry matter; E. sideroxylon: 8 vs. 22 mg·g dry matter), high concentrations of FPCs ( E. melliodora: 5.4 vs. 0.3 mg·g dry matter; E. sideroxylon: 9.8 vs. 0.2 mg·g dry matter) but similar concentrations of nitrogen ( E. melliodora: 15.4 vs. 16.8 mg·g dry matter; E. sideroxylon: 13.1 vs. 14.0 mg·g dry matter). The only difference between the two mosaic trees was in the levels of monoterpenes. The R leaves from the mosaic E. melliodora contained higher concentrations of monoterpenes compared to the S leaves (12 vs. 6 mg·g dry matter). In contrast, the leaves from the E. sideroxylon mosaic contained much higher concentrations of monoterpenes with a reversed pattern (R: 26 vs. S: 45 mg·g dry matter). There were qualitative differences too on the mosaic trees. The R leaves of both species contained much higher concentrations of the monoterpene, 1,8-cineole, whereas the S chemotype of E. sideroxylon only had high concentrations of phellandrenes. Furthermore, the chemical differences between leaves on the R and S branches of the mosaic trees resembled those between the leaves of R and S con-specific trees in the same population. We use these data and knowledge of secondary metabolite biosynthesis to propose that high-level transcriptional differences may control the profile of specialized metabolites in eucalypts. [ABSTRACT FROM AUTHOR]

Summary [ABSTRACT FROM AUTHOR]

Ecologists trying to understand the value of habitat to animals must first describe the value of resources contained in the habitat to animals and, second, they must describe spatial variation in resource quality at a resolution relevant to individual animal foraging. We addressed these issues in a study of koalas (Phascolarcios cinereus) in a Eucalyptus woodland. We measured beneficial and deterrent chemical characteristics as well as the palatability of trees using a near-infrared spectroscopic model based on direct feeding experiments. Tree use. by koalas was influenced by tree size and foliar quality but was also context-dependent: trees were more likely to be visited if they were surrounded by small, unpalatable trees or by large, palatable trees. Spatial autocorrelation analysis and several mapping approaches demonstrated that foliar quality is spatially structured in the woodland at a scale relevant to foraging decisions by koalas and that the spatial structure is an important component of habitat quality. [ABSTRACT FROM AUTHOR]

• Leaves of eucalypt species contain a variety of plant secondary metabolites, including terpenoids and formylated phloroglucinol compounds (FPCs). Both terpene and FPC concentrations are quantitative traits that can show large variation within a population and have been shown to be heritable. The molecular genetic basis of this variation is currently unknown. • Progeny from a field trial of a three-generation mapping pedigree of Eucalyptus nitens were assayed for terpenes and FPCs. Quantitative trait loci (QTL) analyses were conducted using a map constructed from 296 markers to locate regions of the genome influencing foliar concentrations of these plant secondary compounds. • A large number of significant QTL for 14 traits were located across nine linkage groups, with significant clustering of QTL on linkage groups 7, 8 and 9. As expected, QTL for biosynthetically related compounds commonly colocated, but QTL for unrelated monterpenes and FPCs also mapped closely together. • Colocation of these QTL with mapped candidate genes from the various biosynthetic pathways, and subsequent use of these genes in association mapping, will assist in determining the causes of variation in plant secondary metabolites in eucalypts. [ABSTRACT FROM AUTHOR]

Exploring the spatial distribution of variation in plant secondary metabolites is critical for understanding the evolutionary ecology of biochemical diversity in wild organisms. In the present study, concentrations of foliar sideroxylonal, an important and highly heritable defense chemical of Eucalyptus melliodora, displayed strong, fine-scale spatial autocorrelation. The spatial patterns observed could promote associational effects on herbivore foraging decisions, which may influence the selection pressures exerted on sideroxylonal content. Multiple chemical traits have roles in certain eucalypt-herbivore interactions, and the spatial characteristics of the herbivore foraging environment are therefore determined by these different factors. We used a model of E. melliodora intake by common brushtail possums (Trichosurus vulpecula), based on the combined effects of two chemical traits, to explore this idea and found that the spatial patterns were different to those of sideroxylonal alone. Spatial genetic autocorrelation, examined using microsatellites, was strong and occurred at a fine scale, implying that restricted gene flow might allow genetic patches to respond to selection relatively independently. Local two-dimensional genetic autocorrelation, explored using a new heuristic method, was highly congruent with the pattern of local phenotypic variation observed for sideroxylonal, suggesting that the genetic variance underlying the sideroxylonal variation is similarly structured. Our results suggest that the spatial distribution of genetic and phenotypic variation could influence both the selective pressure imposed by herbivores on eucalypt defenses and the potential of populations to respond to natural selection. Spatial context should be considered in future studies of plant-herbivore interactions. [ABSTRACT FROM AUTHOR]

Plant secondary metabolites (PSMs) offer plants chemical defences against herbivores, and are known to influence intake and diet choice in both insect and mammalian herbivores. However, there is limited knowledge regarding how PSMs influence herbivore foraging decisions. Herbivore foraging decisions, in turn, directly impact on which individual plants, and plant species, are selected for consumption. We took advantage of the natural variation in sideroxylonal concentrations in the foliage of Eucalyptus melliodora (Cunn. ex Schauer) to investigate feeding patterns of a marsupial folivore, the common ringtail possum, Pseudocheirus peregrinus (Boddaert 1785). Foliage, collected from six trees, contained between 0.32 and 12.97 mg g-DM-1 sideroxylonal. With increasing sideroxylonal concentrations, possums decreased their total intake, rate of intake and intake per feeding bout, and increased their cumulative time spent feeding. Possums did not alter their total feeding time, number of feeding bouts or time per feeding bout in response to increasing sideroxylonal concentrations. Results demonstrate important behavioural changes in foraging patterns in response to sideroxylonal. These behavioural changes have important implications, in relation to altered foraging efficiency and potential predation risk, for herbivores foraging in the field. As a result, the spatial distribution of dietary PSMs across a landscape may directly influence herbivore fitness, and ultimately habitat selection of mammalian herbivores. [ABSTRACT FROM AUTHOR]

Many studies quantify total phenolics or total tannins, but understanding the ecological role of polyphenolic secondary metabolites requires at least an understanding of the diversity of phenolic groups present. We used UPLC-MS/MS to measure concentrations of different polyphenol groups - including the four most common tannin groups, the three most common flavonoid groups, and quinic acid derivatives - in foliage from 628 eucalypts from the genera Eucalyptus , Angophora and Corymbia . We also tested for phylogenetic signal in each of the phenolic groups. Many eucalypts contained high concentrations of polyphenols, particularly ellagitannins, which have been relatively poorly studied, but may possess strong oxidative activity. Because the biosynthetic pathways of many phenolic compounds share either precursors or enzymes, we found negative correlations between the concentrations of several of the constituents that we measured, including proanthocyanidins (PAs) and hydrolysable tannins (HTs), HTs and flavonol derivatives, and HTs and quinic acid derivatives. We observed moderate phylogenetic signal in all polyphenol constituents, apart from the concentration of the prodelphinidin subunit of PAs and the mean degree of polymerisation of PAs. These two traits, which have previously been shown to be important in determining plants' protein precipitation capacity, may have evolved under selection, perhaps in response to climate or herbivore pressure. Hence, the signature of evolutionary history appears to have been erased for these traits. This study is an important step in moving away from analysing “totals” to a better understanding of how phylogenetic effects influence phenolic composition, and how this in turn influences ecological processes. [ABSTRACT FROM AUTHOR]

Dietary tannins are ubiquitous in woody plants and may have serious negative effects on herbivores by inducing a loss of dietary protein and producing toxins if they are hydrolyzed in the gut. Many herbivorous mammals counter the negative effects of tannins through tannin-binding salivary proteins (TBSPs) that inactivate tannins by forming insoluble complexes and prevent them from interacting with other more valuable proteins. Howlers are the most folivorous New World primates and ingest foods with varying tannin content. We studied the presence of TBSPs in six wild mantled howlers ( Alouatta palliata mexicana) immediately after capture and in captivity when fed on two diets composed of natural ingredients: a mixture of fruit and leaves or only leaves. Protein concentration was determined in whole saliva samples, followed by gel electrophoresis. We identified two protein bands of 17 and 25 kDa that have tannin-binding capacity. Although the monkeys ate almost twice as much condensed tannins in the leaf diet than in the fruits and leaves diet (7 vs. 4 g/d dry matter) the salivary protein concentration did not differ between the two diets (leaf diet: 3.29 ± SE 0.82 vs. fruit and leaves diet: 3.42 ± SE 0.62 mg/ml) and we found no additional protein bands in response to either diet. We suggest that the continuous expression of TBSPs is part of a dietary strategy that enables howlers to consume diets with variable tannin contents, thus partly explaining their dietary flexibility. Although the importance of salivary proteins to arboreal primates is broadly accepted, to our knowledge this is the first report of TBSPs in any Neotropical primate. [ABSTRACT FROM AUTHOR]

A central goal of nutritional ecology is to understand how variation in food quality limits the persistence of wild animal populations. Habitat suitability for browsing mammals is strongly affected by concentrations of nutrients and plant secondary metabolites (PSMs), but our understanding of this is based mostly on short-term experiments of diet selection involving captive animals. In the wild, browsers forage in biologically, chemically and spatially-complex environments, and foraging decisions in response to varying food quality will be correspondingly complicated. We have identified four steps that must be achieved in order to translate our understanding from laboratory experiments to populations of mammalian browsers: 1) knowing what foods and how much of these wild browsers eat, as well as what they avoid eating; 2) knowing the relevant aspects of plant nutritional and defensive chemistry to measure in a given system and how to measure them; 3) understanding the spatial distribution of nutrients and PSMs in plant communities, the costs they impose on foraging and the effects on animals' distributions; and 4) having appropriate statistical tools to analyse the data. We discuss prospects for each of these prerequisites for extending laboratory studies of nutritional quality, and review recent developments that may offer solutions for field studies. We also provide a synthesis of how to use this nutritional knowledge to link food quality to population regulation in wild mammals and describe examples that have successfully achieved this aim. [ABSTRACT FROM AUTHOR]

The idea that biotic interactions, including herbivory, predation and competition are more intense at lower latitudes is widely accepted and underpins several dominant theories on the latitudinal gradient in biodiversity. Current theory also predicts that the intense biotic interactions at low latitudes will select plants for greater defence against herbivores. We reviewed the literature to provide an assessment of the evidence for and against the hypothesis that herbivory is more intense at lower latitudes, and that plants from low latitudes are better defended than are plants from high latitudes. Only 37% of the 38 latitudinal comparisons of herbivory showed higher herbivory at lower latitudes, and the average effect size in a meta-analysis was not significantly different from zero. Thus, the available data do not support the idea that herbivory is generally more intense in the tropics. Only nine of 56 comparisons found higher chemical defences at lower latitudes, and a meta-analysis showed that overall, chemical defences were significantly higher in plants from higher latitudes. This result is counter to the predictions of much of the literature. A meta-analysis showed no significant effect of latitude on physical defence. A review of the literature on feeding trials and common garden experiments showed that herbivores tend to prefer tissue from high latitudes. This trend could stem from differences in overall defence that were not captured by the metrics used in the literature, but could also result from differences in nutritional quality. The empirical data do not support the widespread view that herbivory is generally more intense at lower latitudes, or that plants from low latitudes are generally better defended than are plants from higher latitudes. These results are counter to the prevailing thought on this topic, and suggest that this field may be ripe for the development of new theory. [ABSTRACT FROM AUTHOR]

All free-living animals must make choices regarding which foods to eat, with the choices influencing their health and fitness. An important goal in nutritional ecology is therefore to understand what governs animals’ diet selection. Despite large variation in the availability of different food items, Peruvian spider monkeys ( Ateles chamek) maintain a relatively stable daily protein intake, but allow total energy intake to vary as a function of the composition of available food items. This is referred to as protein-dominated macronutrient balancing. Here we assess the influence of this nutritional strategy on daily and seasonal nutritional intakes, estimate the nutritional value of different foods, and interpret unusual food choices. We conducted continuous all-day observations of focal spider monkeys inhabiting a semideciduous forest in Bolivia. We recorded feeding events, collected foods, and analyzed their nutrient content. By using the Geometric Framework for nutrition, we show that individuals reached their daily end-point in nutrient space —balance between protein and nonprotein energy intake— by consuming nutritionally balanced foods or by alternating between nutritionally complementary foods. The macronutritionally balanced figs of Ficus boliviana were their primary staple food and therefore dominated their overall nutritional intake. Our results also demonstrate that spider monkeys consumed a diverse array of ripe fruits to overcome periods of fig scarcity rather than vice versa; they could obtain sufficient protein on a diet of pure fruit; and unripe figs constituted a nutritionally rewarding and reliable food resource. We hope that the approaches taken and the conclusions reached in this study will catalyze further inquiries into the nutritional ecology of frugivorous primates. [ABSTRACT FROM AUTHOR]

Plant secondary metabolites (PSMs) strongly influence diet selection by mammalian herbivores. Concentrations of PSMs vary within and among plant species, and across landscapes. Therefore, local adaptations may cause different populations of herbivores to differ in their ability to tolerate PSMs. Here, we tested the food intake responses of three populations of a marsupial folivore, the common brushtail possum ( Trichosurus vulpecula Kerr), from different latitudes and habitat types, to four types of PSMs. We found clear variation in the responses of northern and southern Australian possums to PSMs. Brushtail possums from southern Australia showed marked decreases in food intake in response to all four PSMs, while the two populations from northern Australia were not as sensitive and their responses did not differ from one another. These results were unexpected, based on our understanding of the experiences of these populations with PSMs in the wild. Our results suggest that geographically separated populations of possums may have evolved differing abilities to cope with PSMs, as a result of local adaptation to their natural environments. Our results provide the basis for future studies to investigate the mechanisms by which populations of mammalian species differ in their ability to tolerate PSMs. [ABSTRACT FROM AUTHOR]

The concentrations of various foliar chemicals can be estimated by analyzing the spectral reflectance of dried ground leaves. The continuum-removal analysis of Kokaly and Clark [Remote Sens. Environ. 67 (1999) 267] has been an improvement on the standard derivative analysis in such applications. Continuum-removal analysis enables the isolation of absorption features of interest, thus increasing the coefficients of determination and facilitating the identification of more sensible absorption features. The purpose of this study was to test Kokaly and Clark''s methodology with aircraft-acquired hyperspectral data of eucalypt tree canopies, which are more complex than are spectra from many coniferous canopies and much more complex than the spectra from dried ground leaves. The results of the continuum-removal analysis were most encouraging. It identified, in one experiment or another, almost all of the known nitrogen absorption features. The coefficient of determination in one case increased from 0.65, using the standard derivative analysis, to 0.85 with the continuum-removal analysis. It is recommended that continuum-removal analysis become at least a supplement to standard derivative analysis in estimating foliar biochemical concentrations from remote sensing data. This study also reports several other findings: (1) the neural network method generally achieved higher coefficients of determination and lower errors of estimation [root mean square error (RMSE) and standard error of cross validation (SECV)] than did the modified partial least squares (PLS) or stepwise regression methods, probably indicating nonlinear relationships between biochemical concentrations and canopy reflectance; (2) modified partial least squares (MPLS) proved a better statistical method than conventional stepwise regression analysis in many cases in terms of both coefficient of determination and RMSE; and (3) the maximum spectrum of a cluster of tree pixels represents canopy reflectance at least as well as the mean spectrum of the cluster, especially when used in conjunction with the modified partial least squares method. [Copyright &y& Elsevier]