Native plant/herbivore interactions as determinants of the ecological and evolutionary effects of invasive mammalian herbivores: the case of the common brushtail possum

In their native range, mammalian herbivores exist in a suite of direct and indirect ecological and evolutionary relationships with plant populations and communities. Outside their native range these herbivores become embedded in a multitude of new ecological and evolutionary interactions with native plant species in the new range. Sound knowledge of the plant/herbivore interactions in the herbivores’ native range provides an ideal framework to better understand their effects in the introduced range. The example of the common brushtail possum (Trichosurus vulpecula) and its introduction to New Zealand from Australia provides an excellent case study. In Australia, the common brushtail possum is a widespread generalist herbivore and it is thought that this generalist lifestyle has equipped the species well for successful colonisation of New Zealand. In Australia the brushtail possum has co-existed with highly chemically defended foliage since the Oligocene and recent papers have supported the role of possums as agents of selection on eucalypt defences. While the chemical profile of New Zealand foliage is comparatively poorly understood, possums do show clear selectivity between and within populations and some of these interactions may be mediated by the animals ability to ‘cope’ with PSMs, coupled with maintaining its generalist diet. While possums have had less time to effect evolutionary change in New Zealand species, their impacts on plant fitness have been well documented. However, further knowledge on variation and heritability of foliage traits driving possum preferences is needed to elucidate the ecological and evolutionary plant/possum interactions in the invasive range.     

Methodological considerations for the use of faecal nitrogen to assess diet quality in ungulates: The Alpine ibex as a case study

Faecal indices are used to describe spatiotemporal patterns of diet quality in wild herbivores, and near infrared spectroscopy methods (NIRS) distinctively reduce costs and labour compared to conventional chemical analyses. In this study we compared the prediction accuracy of faecal nitrogen (N), ash and fibrous fractions of a laboratory instrument (NIRS) with that of a lightweight portable instrument (VisNIRS) using faecal samples of Alpine ibex (Capra ibex). We then compared how different indices based on faecal N and fibrous fractions were able to detect the expected temporal trend and difference between sexes in diet quality, and verified whether the presumably lower accuracy of NIRS and VisNIRS (compared to wet chemistry) influenced the ability of indices to detect such trends. The laboratory NIRS was more accurate than the portable VisNIRS, especially for fibrous fractions. Predicted data from both instruments clearly identified the temporal trend identified by chemically analysed data, but only NIRS identified the small difference between sexes. Expressing faecal N as a proportion of organic matter (OM) or neutral detergent fibre (NDF) improved the power of the index as compared to expressing it as a proportion of dry matter (DM), irrespective of the method used to estimate the data. Faecal NDF contents followed the patterns expected from the variation of faecal N, while faecal acid detergent fibre (ADF) and especially lignin (ADL) followed contradictory patterns, which might be due to a selection of different forage sources rather than to changing quality of a homogenous forage source. We conclude that expressing faecal N as a proportion of OM is advised with wild herbivores faeces, where ash content can be biased by accidental soil contamination or ingestion. Faecal fibrous fractions are useful to complement faecal N and to avoid simplified interpretations of the results. For this purpose, NIRS methods can be extremely useful to predict multiple indices on large numbers of samples. VisNIRS portable instruments might be unable to detect subtle biological patterns in small numbers of samples. However, developing procedures for portable instruments outside laboratory facilities might further improve practicability, cost effectiveness, and, through allowing on-field predictions on large numbers of samples, power of statistical testing of diet quality indices.     

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.     

Fecal Nitrogen Concentration as a Nutritional Quality Indicator for European Rabbit Ecological Studies

Measuring the quality of the nutritional resources available to wild herbivores is critical to understanding trophic regulation processes. However, the direct assessment of dietary nutritional characteristics is usually difficult, which hampers monitoring nutritional constraints in natural populations. The feeding ecology of ruminant herbivores has been often assessed by analyzing fecal nitrogen (FN) concentrations, although this method has been less evaluated in other taxa. This study analyzed the suitability of FN as an indicator of ingesta quality in the European rabbit (Oryctolagus cuniculus), which is a keystone lagomorph species in Mediterranean ecosystems and of great conservation interest. Firstly, domestic O. cuniculus were used to evaluate under experimental conditions the accuracy of total FN and the metabolic FN as diet quality indicators of forages with characteristics similar to those available under natural conditions. Secondly, the accuracy of Near-Infrared Spectroscopy (NIRS) to calculate FN was tested using partial least squares regression. Thirdly, a pilot field study was conducted to monitor FN dynamics from wild O. cuniculus in three different habitats during wet and drought periods. A strong association was found between diet type and total FN and metabolic FN (Pseudo-R² ≥ 0.89). It was also found that NIRS calibrations were accurate for depicting nitrogen concentrations (R² > 0.98 between NIRS and chemical results). Finally, the seasonal FN dynamics measured in the field were consistent with current knowledge on vegetation dynamics and forage limitations in the three habitats. The results support the use of NIRS methods and FN indices as a reliable and affordable approach to monitoring the nutritional quality of rabbit habitats. Potential applications include the assessment of the mechanistic relationships between resource limitations and population abundance, e.g., in relation to natural drought cycles and to habitat interventions aimed at reinforcing rabbit populations.     

Acid loads induced by the detoxification of plant secondary metabolites do not limit feeding by common brushtail possums (Trichosurus vulpecula)

We fed common brushtail possums artificial diets containing a buffer and the plant secondary metabolite (PSM), orcinol, to test the hypothesis that organic acids, common products of PSM metabolism, limit feeding by common brushtail possums (Trichosurus vulpecula). We introduced several diets containing orcinol and a buffer (urinary alkalising agent) over a course of three experiments. A diet containing 2% orcinol (wet matter) caused possums to reduce their food intake immediately, but feeding returned to normal 1–2 days later. Even though possums excreted strongly acidic urine (pH 5.1) and had perturbed nitrogen metabolism, they maintained their food intake and body mass until the experiment terminated 9 days after the introduction of orcinol. Possums ate 52% less when the basal diet contained 4% orcinol. As expected, the acid loads caused a change in the composition of urinary nitrogen with possums excreting more ammonium than urea and a large amount of unidentified nitrogenous material. Supplementing the diet containing orcinol with buffer neutralised the metabolic acid load and partly restored normal nitrogen metabolism, but did not restore feeding. Also, animals eating orcinol excreted normal amounts of 3-methylhistidine, indicating no increase in muscle protein catabolism. This suggests that a limitation to the rate of detoxification or toxicosis, rather than acid loads, limits the ingestion of acid-inducing PSMs.     

How do soil nutrients affect within-plant patterns of herbivory in seedlings of Eucalyptus nitens?

This study assessed how the palatability of leaves of different age classes (young, intermediate and older) of Eucalyptus nitens seedlings varied with plant nutrient status, based on captive feeding trials with two mammalian herbivores, red-bellied pademelons (Thylogale billardierii), and common brushtail possums (Trichosurus vulpecula). Seedlings were grown under three nutrient treatments (low, medium and high), and we determined how palatability was related to chemical and physical characteristics of the leaves. Pademelons ate more older leaves than young and intermediate leaves for all treatments. This pattern was best explained by sideroxylonals (formylated phloroglucinol compounds known to deter herbivory by other marsupials), and/or essential oil compounds that were present in lower concentrations in older leaves. In the low-nutrient treatment, possums also ate more of the older leaves. However, in the medium- and high-nutrient treatments, possums ate more intermediate leaves than older leaves and showed a behavioural preference for young leaves (consuming younger leaves first) over intermediate and older leaves, in spite of high levels of sideroxylonals and essential oils. The young leaves did, however, have the highest nitrogen concentration of all the leaf age classes. Thus, either sideroxylonals and essential oils provided little or no deterrent to possums, or the deterrent was outweighed by other factors such as high nitrogen. This study indicates that mammalian herbivores show different levels of relative use and damage to leaf age classes at varying levels of plant nutrient status and, therefore, their impact on plant fitness may vary with environment.     

Whole-body protein turnover reveals the cost of detoxification of secondary metabolites in a vertebrate browser

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 [¹⁵N] 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.     

The chemical composition of montane plants and its influence on the diet of the large mammalian herbivores in the Pare National des Volcans, Rwanda

The diets of five large mammalian herbivores in the Pare National des Volcans, a tropical montane forest, were determined using microhistological analysis of faecal material. Correction factors were used to calculate the biomass intake from the relative areas of leaf cuticle found in the faeces. Dietary nitrogen was calculated from the biomass intake obtained in the faecal analysis. This compared well with faecal nitrogen values corrected for dietary intake using equations from the literature. Both measures indicated that each of the five herbivores was obtaining a nitrogen-rich diet. Preferences for food plants were calculated using Manly's alpha as a measure. Nutrient content and other measures of dietary quality of the plants were determined and a multiple regression analysis used to determine which were important for the dietary intake of each herbivore. Mountain gorillas showed a selection of digestible food plants whilst buffalo selected food plants that were not easily digested. Other than in vitro digestibility, none of the measures of dietary quality correlated with the intake of the herbivores. Dietary overlap between the species was significantly lower than expected overlap values calculated using Monte Carlo analyses of neutral models. It is suggested that nutrients were not limiting the populations of these five herbivores.     

Predicting seasonal and spatial variations in diet quality of Pyrenean chamois (Rupicapra pyrenaica pyrenaica) using near infrared reflectance spectroscopy

Monitoring nutritional status of wildlife is key to understanding the population response to changes in food availability. Several direct and indirect methods have been proposed for such purposes, but faecal nitrogen (FN) is by far the most commonly used indirect non-invasive indicator of free-ranging ruminant diet quality. In this work, two methods were used to estimate the FN content in 291 pellet samples of Pyrenean chamois (Rupicapra pyrenaica pyrenaica) collected during a 2-year period (May 2009 to May 2011) in two chamois populations from the Catalan Pyrenees, Spain: the Dumas LECO analyzer and near-infrared reflectance spectroscopy (NIRS). In order to optimize further FN estimations we performed three different NIRS calibrations: (1) general, across year, season and populations; (2) population-specific; and (3) season-specific, i.e., different periods of vegetation growth. FN ranged from 1.66 % to 2.85 % (dry matter basis) and was higher in the main vegetation growing season than in winter. Concerning NIRS calibration, our study confirmed that FN of Pyrenean chamois can be accurately determined by NIRS, since the general calibration was accurate enough to predict FN. It is concluded that NIRS represents a non-destructive, emission-free and low-cost analytical technique which may reduce the use of conventional laborious methods for estimating FN in long-term wildlife monitoring.     

Nutritional Correlates of Koala Persistence in a Low-Density Population

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.     

Inter-population differences in the tolerance of a marsupial folivore to plant secondary metabolites

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.     

The effect of dietary protein quality on nitrogen isotope discrimination in mammals and birds

We tested the competing hypotheses that (1) nitrogen discrimination in mammals and birds increases with dietary nitrogen concentration or decreasing C:N ratios and, therefore, discrimination will increase with trophic level as carnivores ingest more protein than herbivores and omnivores or (2) nitrogen discrimination increases as dietary protein quality decreases and, therefore, discrimination will decrease with trophic level as carnivores ingest higher quality protein than do herbivores. Discrimination factors were summarized for five major diet groupings and 21 different species of birds and mammals. Discrimination did not differ between mammals and birds and decreased as protein quality (expressed as biological value) increased with trophic level (i.e., herbivores to carnivores). Relationships between discrimination factors and dietary nitrogen concentration or C:N ratios were either the opposite of what was hypothesized or non-significant. Dietary protein quality accounted for 72% of the variation in discrimination factors across diet groupings. We concluded that protein quality established the baseline for discrimination between dietary groupings, while other variables, such as dietary protein intake relative to animal requirements, created within-group variation. We caution about the care needed in developing studies to understand variation in discrimination and subsequently applying those discrimination factors to estimate assimilated diets of wild animals.     

The effect of excesses and deficiencies in amino acids on the feeding behaviour of the common brushtail possum (Trichosurus vulpecula)

In this study of the amino acid nutrition of a marsupial we tested three hypotheses: (a) that brushtail possums eat less when diets contain excesses or deficiencies in essential amino acids, (b) that brushtail possums choose diets that do not contain amino acid excesses, and (c) that amino acid consumption is mediated partly by the 5HT3 receptor. Possums ate less when 0.2-1.0% methionine (wet matter) was added to the diet, but similar concentrations of lysine and threonine had little effect. However, when given a choice, possums always selected the basal ration over one with added lysine, methionine or threonine at concentrations between 0.05% and 0.9%. In contrast to the experiments with excess amino acids, possums did not eat less of a diet almost devoid of an essential amino acid. Instead, the possums ate less when their diets contained synthetic amino acids rather than similar amounts and proportions of amino acids as casein. Contrary to the third hypothesis, the 5HT3 receptor antagonist, ondansetron, did not affect feeding by possums given a diet containing 0.8% methionine, suggesting that post-ingestive feedback, via the 5HT3 receptor, does not regulate amino acid intake when diets contain amino acid excesses.     

Predicting Summer Site Occupancy for an Invasive Species,the Common Brushtail Possum (Trichosurus vulpecula), in an Urban Environment

Invasive species are often favoured in fragmented, highly-modified, human-dominated landscapes such as urban areas. Because successful invasive urban adapters can occupy habitat that is quite different from that in their original range, effective management programmes for invasive species in urban areas require an understanding of distribution, habitat and resource requirements at a local scale that is tailored to the fine-scale heterogeneity typical of urban landscapes. The common brushtail possum (Trichosurus vulpecula) is one of New Zealand’s most destructive invasive pest species. As brushtail possums traditionally occupy forest habitat, control in New Zealand has focussed on rural and forest habitats, and forest fragments in cities. However, as successful urban adapters, possums may be occupying a wider range of habitats. Here we use site occupancy methods to determine the distribution of brushtail possums across five distinguishable urban habitat types during summer, which is when possums have the greatest impacts on breeding birds. We collected data on possum presence/absence and habitat characteristics, including possible sources of supplementary food (fruit trees, vegetable gardens, compost heaps), and the availability of forest fragments from 150 survey locations. Predictive distribution models constructed using the programme PRESENCE revealed that while occupancy rates were highest in forest fragments, possums were still present across a large proportion of residential habitat with occupancy decreasing as housing density increased and green cover decreased. The presence of supplementary food sources was important in predicting possum occupancy, which may reflect the high nutritional value of these food types. Additionally, occupancy decreased as the proportion of forest fragment decreased, indicating the importance of forest fragments in determining possum distribution. Control operations to protect native birds from possum predation in cities should include well-vegetated residential areas; these modified habitats not only support possums but provide a source for reinvasion of fragments.     

Roles of the volatile terpene, 1,8-cineole,in plant–herbivore interactions: a foraging odor cue as well as a toxin?

Olfaction is an important sense for many animals, yet its role in foraging by herbivores is poorly known. Many plants contain volatile compounds, such as terpenes, that are not only volatile but can be toxic if ingested. Volatile terpenes can be used by herbivores to assess leaf quality, but there is little evidence for whether they are also used as a searching cue. We applied the giving-up density (GUD) framework to examine fine-scale foraging by two free-ranging mammalian herbivores, the brush-tail possum (Trichosurus vulpecula) and the swamp wallaby (Wallabia bicolor), using patches with food and an inedible matrix that varied in content of a volatile terpene, 1,8-cineole. We tested the effect of (1) increasing dietary cineole concentration, and (2) masking the food odor by adding cineole to the inedible matrix, thus overriding the smell released by the diet. In both species GUD was affected by dietary cineole; a high cineole concentration raised GUD, consistent with its role as a toxin. There was a significant effect of masking on GUD for wallabies but not for possums, suggesting that odor was an important foraging cue at the feeding patch only for the former. Differences in ecological niche and diet may explain this pattern. We suggest that herbivores, such as the swamp wallaby, opportunistically eavesdrop on plant volatiles, i.e., take advantage of the signal proffered for a different function. The cost of this eavesdropping for plants, however, is presumably counteracted by other ecological benefits of these volatiles, including a reduction in leaf consumption as a function of toxicity.     

Effects of tannins on digestion in the common ringtail possum (Pseudocheirus peregrinus), a specialized marsupial folivore

Digestibility trials using common ringtail possums ( Pseudocheirus peregrinus ), small folivorous marsupials, were used to determine effects of tannins on a herbivore which has a specialized hind gut and which normally consumes tannin-rich eucalypt leaves. In one group, untreated leaves were fed to possums, and in a second group polyethylene glycol (PEG) was added to the eucalypt leaves to inactivate tannins. Animals maintained weight, and intakes were not different between the two dietary groups. Digestibility was determined chemically. Digesta were examined histologically. The presence of bacterial rafts in the stomachs of possums fed either diet indicated that PEG did not alter the coprophagic behaviour of possums. Cell wall digestion did not appear to be inhibited by tannins. Tanning of cell contents was predominant, rather than tanning of cell walls. PEG increased faecal excretion of tannins, suggesting that some tannins from normal leaves were digested during gut passage. Dry matter digestibility was higher when animals were fed normal tannin-rich leaves than when they were fed leaves in which tannins were inactivated. The difference could be explained in part by the high digestibility of tannins. The presence of tannin did not reduce nitrogen digestion. We suggest that dissociation of tannin-protein complexes may take place in the specialized caecum. We propose that some other folivorous marsupials may have a similar capacity to overcome tannins. This capacity may allow them to consume a tannin-rich diet.     

Not all types of host contacts are equal when it comes to E. coli transmission

The specific processes that facilitate pathogen transmission are poorly understood, particularly for wild animal populations. A major impediment for investigating transmission pathways is the need for simultaneous information on host contacts and pathogen transfer. In this study, we used commensal Escherichia coli strains as a model system for gastrointestinal pathogens. We combined strain‐sharing information with detailed host contact data to investigate transmission routes in mountain brushtail possums. Despite E. coli being transmitted via the faecal‐oral route, we revealed that, strain‐sharing among possums was better explained by host contacts than spatial proximity. Furthermore, and unexpectedly, strain‐sharing was more strongly associated with the duration of brief nocturnal associations than day‐long den‐sharing. Thus, the most cryptic and difficult associations to measure were the most relevant connections for the transmission of this symbiont. We predict that future studies that employ similar approaches will reveal the importance of previously overlooked associations as key transmission pathways.     

Nitrogen content, amino acid composition and digestibility of fungi from a nutritional perspective in animal mycophagy

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.     

The role of neuropeptides in caterpillar nutritional ecology

Plant diet strongly impacts the fitness of insect herbivores. Immediately, we think of plant defensive compounds that may act as feeding deterrents or toxins. We are, probably, less aware that plants also influence insect growth and fecundity through their nutritional quality. However, most herbivores respond to their environment and select the diet which optimizes their growth and development. This regulation of nutritional balance may occur on many levels: through selecting and ingesting appropriate plant tissue and nutrient digestion, absorption and utilization. Here, we review evidence of how nutritional requirements, particularly leaf protein to digestible carbohydrate ratios, affect caterpillar herbivores. We propose a model where midgut endocrine cells assess and integrate hemolymph nutritional status and gut content and release peptides which influence digestive processes. Understanding the effects of diet on the insect herbivore is essential for the rational design and implementation of sustainable pest management practices.     

哺乳动物的消化策略（英文）

理解动物的营养生态位是充分理解其整个生态学的基础,对于害兽控制和物种保护也很重要,食肉动物的小肠很发达,这可能与对食物的高消化能力有关;杂食性动物有更复杂的胃肠器官,其后端有可进行发酵的盲肠,消化物的平均滞留时间（mean retention times,MRTs)更长;最长的平均滞留时间见于食草动物,其消化道内高密度的微生物种群对不同滞留区内的消化物进行发酵,但是,并不是所有的食草动物都能够最大程度地消化植物纤维,只有反刍动物、骆驼和个体较大的后肠发酵动物（hindgut fermenter)能够具有这种能力,对比而言,许多其它的食草动物,如前肠发酵的有袋类和小型的后肠发酵动物如兔子、田鼠和负鼠等,它们具备可以使植物纤维消化效率最大的消化系统,可以在食物中的纤维素含量非常高的情况下仍能处理大量的食物。这些不同的消化策略使哺乳动物具有广幅的营养生态位。