Optimal digestive strategies for arboreal herbivorous mammals in contrasting forest types: Why Koalas and Colobines are different

Abstract The eucalypt forests of temperate eastern Australia contrast with many forests in tropical Australia, Asia, Africa and Central America in terms of degrees of sclerophylly, concentrations of nutrients in foliage, types and amounts of chemical challenges posed for herbivores, and the range of alternatives to foliage as food. Comparisons between digestive strategies employed by arboreal herbivorous marsupials and primates inhabiting these forests reveal several trends: (i) foregut-fermenters occur only in tropical and sub-tropical forests, whereas caecum and caecum-colon fermenters occur in both temperate eucalypt and the tropical forests; (ii) the arboreal marsupials and primates (koalas, greater gliders, ringtail possums and sportive lemurs) that are the most folivorous are caecum or caecum-colon fermenters with colonic specializations for selective retention of small panicles of digesta; (iii) no caecum or caecum- colon fermenters lacking colonic separation mechanisms achieve consistently high degrees of folivory; (iv) even the most folivorous foregut-fermenting arboreal marsupials (tree kangaroos) or primates (colobus monkeys) include a substantial proportion of high-quality components (fruits or seeds) in their diet. Caecum or caecum-colon fermentation with colonic separation is postulated to be essential for a high degree of folivory in small mammals because it offsets potential limits on intake of high-fibre food and minimizes losses of faecal nitrogen that otherwise would be prohibitive. This digestive strategy is common among folivores in eucalypt forests because the low abundance of large seeds and fleshy fruits limits folivory-frugivory and folivory-granivory. Foregut fermentation employing a sacciform/ tubiform forestomach (as in tree kangaroos and colobus monkeys) is postulated to be optimal for mixed leaf/fruit and leaf/seed diets that are available in many tropical forests, assuming it allows high-fibre meals to be retained for microbial fermentation while permitting low-fibre meals to pass rapidly to the hindstomach for acid/enzymic digestion. However, foregut fermentation appears innappropriate for either primary frugivory or exclusive folivory due to the inefficiency of microbial digestion of simple sugars and constraints on microbial fermentation as a primary energy source for small mammals.     

A simple, integrative assay to quantify nutritional quality of browses for herbivores

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. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Comparative dietary ecology of two congeneric marsupial folivores

Knowledge of species diets is critical to assisting our understanding of their ecology. Using microhistological analysis of faecal samples, we described and compared the diets of sympatrically occurring folivorous congenerics, common and mountain brushtail possums (Trichosurus vulpecula and T. cunninghami, respectively). Throughout the 28‐month study period, common brushtails relied heavily on eucalypt foliage, particularly very young leaves, which is consistent with data from captive studies on their dietary physiology. In contrast, eucalypt foliage formed only a small part of the diet of mountain brushtails, which instead relied heavily on silver wattle foliage. The mean number of plant groups per faecal sample was significantly greater for common brushtails than mountain brushtails. No significant differences in diet between male and female mountain brushtails were detected. However, intraspecific differences in diet occurred for common brushtails: the diet of females included significantly less eucalypt foliage and significantly more foliage of the exotic, tree lucerne, than that of males during the Wet Season (April–November), but not during the Dry Season (December–March). Diets varied temporally for both species, with some individuals feeding on seasonally available resources. The diets described for common and mountain brushtails are consistent with those of a dry‐adapted and mesic‐adapted species, respectively. We discuss how our results contribute to our understanding of the evolutionary history of both study species, and more broadly the family Phalangeridae to which they belong. We also consider the diet of our study species in the context of recent advances in our understanding of interactions between plants that produce secondary metabolites, and mammals specialized to feed on them.     

A comparative study of the effect of two-stage olive cake added to alfalfa on digestion and nitrogen losses in sheep and goats

This work was conducted to evaluate the effect of tannins, contained in a by-product derived from olive oil extraction (two-stage olive cake), on nutrient digestibility, nitrogen (N) losses and liver and kidney functions in goats and wethers. Six Segureña wethers and six Granadina goats were fed three experimental diets: alfalfa hay (AH); alfalfa hay and a concentrate (formulated with two-stage olive cake, barley and a mineral-vitamin mixture) without (AHCO) or with polyethylene glycol (PEG) addition (AHCOP). The inclusion of two-stage olive cake increased condensed tannins content of the diet and led to a decrease ( P < 0.001) in dry matter, organic matter, neutral-detergent fibre, acid-detergent fibre and crude protein (CP) digestibility. PEG supply increased ( P < 0.001) CP digestibility and N losses in urine and faeces only in wethers. Concentrations of creatinine and alkaline phosphatase activity, measured in the serum of animals, were not modified either by olive cake inclusion or by PEG supply. Results from this work indicate a higher sensitivity of sheep to diets containing condensed tannins, compared with goats, and reveal no toxic effect of tannins from two-stage olive cake at the concentration used in the experimental diets.     

Lotus corniculatus condensed tannins decrease in vivo populations of proteolytic bacteria and affect nitrogen metabolism in the rumen of sheep

Condensed tannins in forage legumes improve the nutrition of sheep by reducing ruminal degradation of plant protein and increasing crude protein flow to the intestine. However, the effects of condensed tannins in forage legumes on rumen bacterial populations in vivo are poorly understood. The aim of this study was to investigate the specific effects of condensed tannins from Lotus corniculatus on four proteolytic rumen bacteria in sheep during and after transition from a ryegrass (Lolium perenne)-white clover (Trifolium repens) diet (i.e., low condensed tannins) to a Lotus corniculatus diet (i.e., higher condensed tannins). The bacterial populations were quantified using a competitive polymerase chain reaction. Lotus corniculatus was fed with or without ruminal infusions of polyethylene glycol (PEG), which binds to and inactivates condensed tannins, enabling the effect of condensed tannins on bacterial populations to be examined. When sheep fed on ryegrass-white clover, populations of Clostridium proteoclasticum B316T, Butyrivibrio fibrisolvens C211a, Eubacterium sp. C12b, and Streptococcus bovis B315 were 1.5 x 10(8), 1.1 x 10(6), 4.6 x 10(8), and 7.1 x 10(6) mL(-1), respectively. When the diet was changed to Lotus corniculatus, the average populations (after 8-120 h) of C. proteoclasticum, B. fibrisolvens, Eubacterium sp., and S. bovis decreased (P < 0.001) to 2.4 x 10(7), 1.1 x 10(5), 1.1 x 10(8), and 2.5 x 10(5) mL(-1), respectively. When PEG was infused into the rumen of sheep fed Lotus corniculatus, the populations of C. proteoclasticum, B. fibrisolvens, Eubacterium sp., and S. bovis were higher (P < 0.01-0.001) than in sheep fed Lotus corniculatus without the PEG infusion, with average populations (after 8-120 h) of 4.9 x 10(7), 3.8 x 10(5), 1.9 x 10(8), and 1.0 x 10(6), respectively. Sheep fed the Lotus corniculatus diet had lower rumen proteinase activity, ammonia, and soluble nitrogen (P < 0.05-0.001) than sheep that were fed Lotus corniculatus plus PEG. The Lotus corniculatus diet reduced rumen nitrogen digestibility (P < 0.05) and ammonia pool size and increased the flow of undegraded feed nitrogen to the abomasum. The nitrogen intake, rumen non-ammonia nitrogen pool size, rumen microbial non-ammonia nitrogen pool size, and abomasal microbial non-ammonia nitrogen fluxes were similar both in sheep fed only Lotus corniculatus and in sheep fed Lotus corniculatus plus PEG, but nonmicrobial non-ammonia nitrogen flux to the abomasum was higher (P < 0.01) for the sheep fed only Lotus corniculatus. Although condensed tannins in Lotus corniculatus reduced the populations of some proteolytic bacteria, total ruminal microbial protein and microbial protein outflow to the abomasum were unchanged, suggesting a species-specific effect of condensed tannins on bacteria in the rumen.     

Diet selection in the green ringtail possum (Pseudochirops archeri): A specialist folivore in a diverse forest

Abstract This study investigated the feeding ecology of the green ringtail possum, Pseudochirops archeri (Pseudocheiridae) in a tropical rainforest with 94 plant species in the canopy. Over 50% of tree use was from only four tree species, Aleurites rockinghamensis, Ficus fraseri, Arytera divaricata and Ficus copiosa. These species were used significantly more frequently than would be expected if tree species were selected randomly in proportion to their relative abundance in the forest. Conversely, 88 other tree species present were used less frequently than expected. Possums also favoured particular individual trees within some of the preferred tree species. In 91% of feeding observations, possums consumed mature leaves only. The availability of young leaves, flowers and fruit varied throughout the year, with a peak in availability of these resources during the early wet season. By primarily selecting mature leaves, green ringtail possums reduce their dependence on seasonally variable resources. We suggest that green ringtail possums should be considered as specialist folivores, focusing on only a few of the tree species available, possibly due to advantages associated with limiting the number of plant secondary metabolites in their diet. Furthermore, they favour certain individual trees within species, perhaps due to intraspecific variation in plant secondary metabolites or nutrient content, behaviour that has been well established in eucalypt folivores. We conclude that green ringtail possums are highly specialized in their feeding ecology, limiting their diet to a small number of continuously available food items.     

What physiological processes permit insects to eat Eucalyptus leaves?

Many species of insects eat Eucalyptus foliage despite its relatively low nutritional value and the many plant secondary metabolites (PSMs) present, for example, terpenes, phenols and formylated phloroglucinols (FPGs). Formylated phloroglucinols are a new class of PSMs that act as antifeedants for possums and koalas. What physiological processes are present that permit insects to eat eucalypt foliage and how do PSMs influence insect feeding or digestion? Some trees seem to be repeatedly infested with eucalypt‐feeding insects, possibly as a result of previous chemosensory cues remaining from parental selection of a plant. Avoidance or storage of PSMs permit jarrah leafminers (Perthida glyphopa) and sawflies (Perga sp.) to consume eucalypt foliage without dealing with the majority of these compounds. Some PSMs can be metabolized by polysubstrate membrane oxidases as found in caterpillars or sawflies that feed on eucalypts. High midgut pH may be advantageous for nutrient extraction and PSM metabolism, and midgut pH ranges between 8.5 and 8.9 for caterpillars of Hyalarcta huebneri. Plant secondary metabolites may not be absorbed as a result of the combined presence of the peritrophic matrix and endogenous surfactants. Excretion of PSMs can be as metabolites or intact compounds. Both putative metabolites and sideroxylonal‐A, an FPG, are present in the faeces of larvae of the case moth, H. huebneri. The presence of sideroxylonal‐A in the food had an effect on the presence of 5‐hydroxytryptamine (5HT) in the central nervous system of caterpillars, as larvae fed leaves with a high concentration of sideroxylonal‐A had relatively more 5HT in the brain and central nervous system ganglia than larvae fed leaves containing a low concentration. Further work is necessary to clarify how PSMs are handled by eucalypt‐feeding insects and what effect FPGs have on feeding and digestion.     

Differential susceptibility to Eucalyptus secondary compounds explains feeding by the common ringtail (Pseudocheirus peregrinus) and common brushtail possum (Trichosurus vulpecula)

The effect of two plant secondary metabolites, tannins and formylated phloroglucinol compounds (FPCs), on the intake of Eucalyptus foliage by common ringtail (Pseudocheirus peregrinus) and common brushtail possums (Trichosurus vulpecula) was studied. We manipulated the amount of tannin that was free to bind with protein by coating foliage with polyethylene glycol 4000 (PEG) and relied on natural intraspecific variation in FPC concentrations. In contrast to ringtail possums, brushtail possums showed a greater tolerance to FPCs and ate more foliage when it was coated with PEG, suggesting that tannins limited their food intake. Brushtails detected the effects of tannins through immediate oral sensations rather than through systemic effects. Ringtail possums appeared highly tolerant of foliar tannins yet susceptible to low concentrations of FPCs. We could not detect any interaction between tannins and FPCs that affected the intake of Eucalyptus foliage by either species of possum. Although ringtail and brushtail possums are widely regarded as specialist and generalist folivores, respectively, their differential susceptibility to co-occurring secondary metabolites suggests greater complexity. Each possum species appears to be a specialist in its own right, which leads to a partitioning of available foliage. Brushtails avoid tannins and ringtails avoid FPCs.     

Digestive strategies and food choice in mantled howler monkeys Alouatta palliata mexicana: bases of their dietary flexibility

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.     

Gut pH, redox conditions and oxygen levels in an aquatic caterpillar: potential effects on the fate of ingested tannins

Larvae of the freshwater moth Acentria ephemerella (Pyralidae, Lepidoptera) can fully develop on Myriophyllum spicatum, a submerged macrophyte containing 7-10% of its dry mass as tannins. We investigated the physicochemical gut parameters of larvae fed with M. spicatum or Potamogeton perfoliatus, a food plant lacking tannins, and the chemical fate of ingested polyphenols. Microelectrode studies revealed that larval midguts were slightly alkaline (pH 8) and had a positive redox potential. Whole guts were oxygen sinks owing to the oxygen demand of the gut contents. Oxygen penetrated the midgut up to 100 microm, but the centres of the foregut and midgut were always anoxic. The physicochemical parameters of the guts did not change with the food plant. The major tannin from M. spicatum, tellimagrandin II, was significantly depleted in the midgut and was not detected in faeces. In vitro studies indicated that tellimagrandin II is rapidly depleted mainly through oxidation, and hydrolysis might also occur. Our findings for A. ephemerella are compared with those for terrestrial Lepidoptera, and possible mechanisms for adaptations to tannin-rich food plants are discussed.     

Effects of dietary quality on basal metabolic rate and internal morphology of European starlings (Sturnus vulgaris)

European starlings (Sturnus vulgaris) were fed either a low- or high-quality diet to test the effects of dietary quality on basal metabolic rate (BMR) and internal morphology. Basal metabolic rate did not differ significantly between the two dietary groups, but internal morphology differed greatly. Starlings fed the low-quality diet had heavier gastrointestinal tracts, gizzards, and livers. Starlings fed the high-quality diet had heavier breast muscles. Starlings on the low-quality diet maintained mass, while starlings on the high-quality diet gained mass. Dry matter digestibility and energy digestibility were lower for starlings fed the low-quality diet, and their food and water intake were greater than starlings on the high-quality diet. The lack of dietary effect on BMR may be the result of increased energy expenditure of digestive organs paralleling a reduction of energy expenditure of organs and tissues not related to digestion (i.e., skeletal muscle). This trade-off in energy allocation among organs suggests a mechanism by which organisms may alter BMR in response to a change in seasonal variation in food availability.     

A pilot study on the ontogeny of digestive physiology in Japanese macaques (Macaca fuscata)

An assumption based on the Jarman–Bell principle suggests a positive relationship between body size and the digestive efficiency in animals, where smaller animals are less effective at digesting fibrous food due to shorter digesta passage. To examine the effect of body size within a species and explore a potential physiological background of ontogenetic diet shifts, we measured food intake, digestibility, digesta passage and gut fill in nine Japanese macaques, including three juveniles/subadult animals. Although these three showed a comparable digestive efficiency as the older animals on a low-fiber diet, they did not achieve the long retention times of adults in spite of similar levels of indigestible food intake and gut capacity. While the limited sample size would not allow generalized conclusions on ontogenetic digestive development in primates, this study suggests additional, yet unexplored effects other than food intake, digestion and gut capacity on digesta retention during ontogeny.     

Bactericidal effect of hydrolysable and condensed tannin extracts on <Emphasis Type="Italic">Campylobacter jejuni</Emphasis> in vitro

Strategies are sought to reduce intestinal colonisation of food-producing animals by Campylobacter jejuni, a leading bacterial cause of human foodborne illness worldwide. Presently, we tested the antimicrobial activity of hydrolysable-rich blackberry, cranberry and chestnut tannin extracts and condensed tannin-rich mimosa, quebracho and sorghum tannins (each at 100 mg/mL) against C. jejuni via disc diffusion assay in the presence of supplemental casamino acids. We found that when compared to non-tannin-treated controls, all tested tannins inhibited the growth of C. jejuni and that inhibition by the condensed tannin-rich mimosa and quebracho extracts was mitigated in nutrient-limited medium supplemented with casamino acids. When tested in broth culture, both chestnut and mimosa extracts inhibited growth of C. jejuni and this inhibition was much greater in nutrient-limited than in full-strength medium. Consistent with observations from the disc diffusion assay, the inhibitory activity of the condensed tannin-rich mimosa extracts but not the hydrolysable tannin-rich chestnut extracts was mitigated by casamino acid supplementation to the nutrient-limited medium, likely because the added amino acids saturated the binding potential of the condensed tannins. These results demonstrate the antimicrobial activity of various hydrolysable and condensed tannin-rich extracts against C. jejuni and reveal that condensed tannins may be less efficient than hydrolysable tannins in controlling C. jejuni in gut environments containing high concentrations of amino acids and soluble proteins.     

Effect of tree leaves with or without urea as a feed supplement on nutrient digestion and nitrogen balance in sheep

Two nitrogen balance experiments in a 4 × 4 Latin square design were conducted to compare leaves of Grewia oppositifolia and Ziziphus mauritiana containing condensed tannins (CT) 0.08 g and 34.7 g per kg DM, respectively as supplement on feed intake, nutrients digestibility and nitrogen (N) retention in adult wethers and to examine the benefits of adding urea to Z. mauritiana leaves containing higher level of CT on N utilization in the animals. In experiment 1, the basal diet of oat hay was fed to sheep either un-supplemented or supplemented with 320 g dried leaves of Z. mauritiana or 320 g dried leaves of G. oppositifolia or 160 g dried leaves each of Z. mauritiana and G. oppositifolia. In vivo dry matter (DM) digestibility was not different among the four diets while N digestibility remained significantly lower (P<0.05) in Z. mauritiana supplemented diets. Daily intake of oat hay reduced (P<0.05) and that of total diet DM increased (P<0.001) with feeding of the leave supplements. Mean N retention on control diet was 4.39 g/d and increased to 7.51 g/d with inclusion of G. oppositifolia leaves but did not change with the other two supplements. In experiment 2, a basal diet of sorghum hay was fed ad libitum without any supplement (control diet) and the other three diets were supplemented with 320 g dried leaves of Z. mauritiana with no urea, 5 g urea/d or 10 g urea/d. Daily intake (g/d) of sorghum hay or total DM intake did not respond to inclusion of urea in the diets. Supplementation of Z. mauritiana with or without urea did not affect digestibility of DM, organic matter and acid detergent fiber. Nitrogen retention increased (P<0.05) from 0.57 g/d on control diet to 3.72 g/d with supplementation of Z. mauritiana leaves. Addition of urea 5 g/d did not further increase the N retention (4.78 g/d) but was significantly increased to 7.16 g/d in response to 10 g urea/d in the diet. It was concluded that response to urea feeding in the presence of tannin rich Z. mauritiana leaves was dose-dependant and that feeding 10 g urea/d increased the capacity of sheep to consume more feed and retain more N in the body.     

Digestive plasticity and the cost of acclimation to dietary chemistry in the omnivorous leaf-eared mouse Phyllotis darwini

We examined the costs associated with acclimation in an omnivorous mouse (Phyllotis darwini) fed two contrasting diets (carbohydrate-rich and protein-rich). We studied the response of gut morphology and digestive performance in animals shifted to a novel diet at different developmental stages. When acclimated adult animals were shifted to the alternative diet, energy digestibility decreased. We also found long-term consequences to diet acclimation. Animals reacclimated for 15 days to an alternative diet did not increase digestive performance. Although no effects of diet on gut morphology were noted, a significant positive correlation between energy digestibility and small intestinal length was found, explaining most of the variability observed in energy digestibility. These results suggest that caution should be used when defining adaptive changes if the possible cost of acclimation is neglected. Accepted: 18 May 2000     

Effects of Acomastylis rossii tannins on a mammalian herbivore,the North American pika,Ochotona princeps

 I investigated the effects of tannin consumption, using plant tannins naturally occurring in the diet, on a herbivorous mammal, the North American pika, Ochotona princeps. The objectives were to determine if a high-tannin diet influenced protein and dry matter apparent digestibility, fiber digestibility and production of detoxification by-products. Additionally, I examined the possibility that pikas produce salivary tannin-binding proteins, a potential mechanism for avoiding detrimental effects of tannins. My results demonstrate that although pikas constitutively produce salivary tannin-binding proteins, animals consuming a high-tannin diet of Acomastylis rossii exhibited lower dry matter, protein and fiber digestion and excreted higher concentrations of detoxification by-products. Thus, A. rossii tannins are potential toxins as well as digestibility reducers. I propose a hypothesis coupling detoxification to reduced fiber digestion that is applicable to pikas as well as other mammalian herbivores consuming phenolic-rich diets. Received: 31 July 1995 / Accepted: 9 July 1996     

Selective herbivory by brushtail possums: Determining the age of ingested leaves using n‐alkanes

Abstract Leaves often decline in nutritional quality as they age, and selective feeding on young leaves may be nutritionally important for herbivores. Preference by mammalian herbivores for young leaves has rarely been measured in the field owing to technical difficulties. We measured preferences with respect to leaf age of an arboreal folivore, the brushtail possum (Trichosurus vulpecula Kerr), feeding on southern rata (Metrosideros umbellata Cav.; Myrtaceae) in a new application of the alkane technique. We characterized the cuticle waxes (n‐alkanes) of rata leaves that were less than 1 year old (‘1‐year’), between 1 and 2 years (‘2‐year’) and greater than 2 years old (‘>2‐year’). Simulations showed that the method accurately discriminated between 1‐year and other age groups but slightly overestimated the importance of rare components of the diet. This bias was larger when discriminating between 2‐year and >2‐year leaves apparently because they had more‐similar alkane profiles. Metrosideros umbellata leaf formed 20.8% of the diet of a population of possums from Rakiura, New Zealand, sampled in autumn 2002 (n = 33). Of the M. umbellata component, alkane analyses showed that 1‐year leaves formed 88.7 ± 3.9% of the diet despite making up only 39.5 ± 2.2% of the leaf biomass on rata trees (n = 14). The foliar concentrations of the macronutrients N, P and K all declined significantly with leaf age (P < 0.0001). Lignin content did not measurably increase with leaf age, suggesting that digestibility per se did not determine the preference of brushtail possums for young rata leaves. This study provides the first quantitative evidence that possums discriminate by leaf age and that the resulting diet is enriched in macronutrients.     

Food selection by the silver leaf monkey,Trachypithecus auratus sondaicus,in relation to plant chemistry

Summary Samples of leaves and fruits exploited as food items byTrachypithecus auratus sondaicus were analysed for nitrogen content, acid detergent fibre (ADF), pepsin cellulase digestibility (CDIG), condensed tannins (CT), total phenolics (TP) and protein precipitation capacity (PP) and compared with fruits and leaves not eaten. Differences in chemical measures for items eaten and not eaten were not statistically significant but trends indicate that leaves may have been selected for their lower fibre content and greater digestibility. Fruits eaten also had a higher mean level of CDIG and lower mean level of ADF than fruits not eaten but these measures are not considered to be of major importance in fruit selection as CDIG was lower and ADF higher in fruits eaten than in leaves eaten. Levels of CT, TP and PP capacity were higher in fruits eaten than in fruits not eaten but lower in leaves eaten than in leaves not eaten. The role of tannins and phenolics in food selection is discussed. Leaves (and fruits) were not strongly selected on the basis of protein content. Approximately half the dietary intake ofT. auratus sondaicus was leaves, a protein-rich food source. Possibly, protein levels in foliage at Pangandaran were sufficient that selection for this nutrient was not required. A nutrient other than protein (for example, soluble carbohydrates) may have been maximised through food selection. The protein/ADF ratio may provide an indicator of the acceptability of foliage in a habitat as potential food for a colobine. However, this ratio did not govern food selection byT. auratus sondaicus at Pangandaran.     

African rainforest vegetation and rumen microbes: Phenolic compounds and nutrients as correlates of digestibility

Summary In order to refine hypotheses concerning food selection by generalist herbivores with ruminant-like digestive systems the chemical correlates of digestibility in such a system have been studied. Samples of seeds and leaves from tree species growing in two African rainforests (Douala-Edea Forest Reserve, Cameroon, and Kibale Forest, Uganda) were assayed for phenolic content and nutrient content, and in-vitro dry matter digestibility was analysed utilizing rumen inoculum from a fistulated steer. Both forests studied carry populations of colobine monkeys with ruminant like digestive tracts. Content of condensed tannins and, to a lesser extent of total phenolics, was found to be negatively correlated with digestibility; a result that may be attributable to the inctivation of microbial enzymes by tannins. The negative association of tannin content and digestibility was stronger in material from the Cameroon site, the vegetation of which contains considerably higher concentrations of tannins and is generally less digestible than that from the Ugandan site. Gross energy content of leaves was also found to be persistently negatively correlated with digestibility. The interpretation of this result is uncertain; however, gross energy yield may well reflect variation in content of cell wall polymers, especially lignin. For the complete set of data, tannins presented the strongest observed correlation with digestibility, but when only mature leaves were considered the relationship with gross energy appeared stronger. No strong association was noted between high nutrient content and high digestibility. This was attributed to the fact that the assay measured the extent of digestion under standardised and very favourable conditions of nutrient supply. The results obtained are discussed in relation to observations of leaf and seed selection preferences of Colobus spp. in these two forests.Publication 19-013 of the Wisconsin Regional Primate Research Center     

Galagos and Gummivory

An experimental program investigating the role of gum in the diet of captiveGalago senegalensis braccatus is reviewed. One set of studies investigated the role of gum chemistry in dietary preferences. The animals showed a preference forAcacia gums over gums from other plants. If condensed or hydrolized tannins were added to theAcacia gums, such gums were consumed at a lower rate than plainAcacia gum. This effect was stronger with the condensed (quebracho) tannin. The effect of such secondary compounds on gum consumption in wild galagos needs investigation. The second set of studies examined digestion of gums, which are complex polysaccarides probably requiring fermentation in the gut. Substitution of gum for simple carbohydrate sources in the diet lead to increased gut transit time. On the gum diet, animals maintained weight without an increase in intake or a decrease in activity levels. Apparent dry matter digestibility of the two diets was equivalent. Indicators of gut bacterial action increased on the gum diet. These observations are strong indirect evidence for an increase of fermentation in the gut on the gum diet.