The Role of Individual Traits and Environmental Factors for Diet Composition of Sheep

Large herbivore consumption of forage is known to affect vegetation composition and thereby ecosystem functions. It is thus important to understand how diet composition arises as a mixture of individual variation in preferences and environmental drivers of availability, but few studies have quantified both. Based on 10 years of data on diet composition by aid of microhistological analysis for sheep kept at high and low population density, we analysed how both individual traits (sex, age, body mass, litter size) linked to preference and environmental variation (density, climate proxies) linked to forage availability affected proportional intake of herbs (high quality/low availability) and Avenella flexuosa (lower quality/high availability). Environmental factors affecting current forage availability such as population density and seasonal and annual variation in diet had the most marked impact on diet composition. Previous environment of sheep (switch between high and low population density) had no impact on diet, suggesting a comparably minor role of learning for density dependent diet selection. For individual traits, only the difference between lambs and ewes affected proportion of A. flexuosa, while body mass better predicted proportion of herbs in diet. Neither sex, body mass, litter size, ewe age nor mass of ewe affected diet composition of lambs, and there was no effect of age, body mass or litter size on diet composition of ewes. Our study highlights that diet composition arises from a combination of preferences being predicted by lamb and ewes’ age and/or body mass differences, and the immediate environment in terms of population density and proxies for vegetation development.     

Body Condition Indices Predict Reproductive Success but Not Survival in a Sedentary,Tropical Bird

Body condition may predict individual fitness because those in better condition have more resources to allocate towards improving their fitness. However, the hypothesis that condition indices are meaningful proxies for fitness has been questioned. Here, we ask if intraspecific variation in condition indices predicts annual reproductive success and survival. We monitored a population of Neochmia phaeton (crimson finch), a sedentary, tropical passerine, for reproductive success and survival over four breeding seasons, and sampled them for commonly used condition indices: mass adjusted for body size, muscle and fat scores, packed cell volume, hemoglobin concentration, total plasma protein, and heterophil to lymphocyte ratio. Our study population is well suited for this research because individuals forage in common areas and do not hold territories such that variation in condition between individuals is not confounded by differences in habitat quality. Furthermore, we controlled for factors that are known to impact condition indices in our study population (e.g., breeding stage) such that we assessed individual condition relative to others in the same context. Condition indices that reflect energy reserves predicted both the probability of an individual fledging young and the number of young produced that survived to independence, but only during some years. Those that were relatively heavy for their body size produced about three times more independent young compared to light individuals. That energy reserves are a meaningful predictor of reproductive success in a sedentary passerine supports the idea that energy reserves are at least sometimes predictors of fitness. However, hematological indices failed to predict reproductive success and none of the indices predicted survival. Therefore, some but not all condition indices may be informative, but because we found that most indices did not predict any component of fitness, we question the ubiquitous interpretation of condition indices as surrogates for individual quality and fitness.     

Surface enlargement in the rumen of free-ranging muskoxen (<Emphasis Type="Italic">Ovibos moschatus</Emphasis>)

The intraruminal papillation pattern indicates the degree of rumen contents stratification and is related to the feeding niche of a ruminant. Muskoxen (Ovibos moschatus) display a variety of morphophysiological adaptations typical for grazers. We investigated the intraruminal papillation of 22 free-ranging muskoxen from five different months by comparing the surface enlargement factor both between seasons and between individual rumen regions. The seasonal pattern of rumen papillation indicated a distinct seasonality in food quality. The intraruminal papillation indicated a moderate degree of rumen contents stratification typical for intermediate feeders. The nutritional ecology of muskoxen is characterised by specific morphophysiological adaptations to a grass-dominated diet that nevertheless allow extensive seasonal use of browse forage.     

Validating Predictive Models of Nutritional Condition for Mule Deer

Abstract: We developed new, and validated existing, indices of nutritional condition for live and dead mule deer (Odocoileus hemionus). Live animal indices included a body condition score (BCS), thickness of subcutaneous fat and selected muscles using ultrasonography, and body mass. Dead animal indices included femur, metatarsal, and mandible marrow fat, 3 kidney fat indices, and 2 carcass scoring methods. We used 21 female deer and 4 castrates (1-11 yr old) varying widely in nutritional condition (2-28% ingesta-free body fat). Deer were euthanized and homogenized for chemical analysis of fat, protein, water, and ash content. Estimates of fat and gross energy (GE) were regressed against each condition indicator using regression. Subcutaneous fat thickness, a rump BCS, and rLIVINDEX (an arithmetic combination of subcutaneous fat thickness and the rump BCS) were most related to condition for live animals (r² ≥ 0.87, P < 0.001) whereas the Kistner score and kidney fat were most related to fat and GE for dead animals (r² ≥ 0.77, P < 0.001). We also evaluated range of usefulness and sensitivity to small changes in body condition for all models. In general, indices with moderate or highly curvilinear statistical relations to body fat or those based on only one fat depot or a small number of ranking scores will have limitations in their use. Our results identify robust tools for a variety of research and monitoring designs useful for evaluating nutrition's effect on mule deer populations.     

Can Body Condition and Somatic Indices be Used to Evaluate Metal-Induced Stress in Wild Small Mammals?

Wildlife is exposed to natural (e.g., food availability and quality, parasitism) and anthropogenic stressors (e.g., habitat fragmentation, toxicants). Individual variables (e.g., age, gender) affect behaviour and physiology of animals. Together, these parameters can create both great inter-individual variations in health indicators and interpretation difficulties. We investigated the relevance of body condition and somatic indices (liver, kidneys) as indicators of health status in wood mice (Apodemus sylvaticus, n = 560) captured along a metal pollution gradient in four landscape types (30 sampling squares 500-m sided). The indices were calculated using a recently proposed standard major axis regression instead of an ordinary least square regression. After considering age and gender for the body condition index, no landscape type influence was detected in the indices. However, important index variability was observed between sampling squares; this effect was included as a random effect in linear models. After integrating all individual and environmental variables that may affect the indices, cadmium (Cd) concentrations in both the liver and kidneys were negatively related to body condition and liver indices only for individuals from highly contaminated sites. Lead in the liver was negatively related to the liver index, and Cd in kidneys was positively linked to the kidney index, potentially suggesting metal-induced stress. However, interpretation of these indices as a wildlife ecotoxicology tool should be performed with caution due to the sensitivity of potentially confounding variables (e.g., individual factors and environmental parameters).     

Significance of diet type and diet quality for ecological diversity of African ungulates

1. We test two nutritional hypotheses for the ecological diversity of ungulates, the browser/grazer (diet type) and diet quality models, among free-ranging herbivores in a South African savanna, the Kruger National Park. Tests are based on assessment of relationships between diet type and diet quality with body mass and hypsodonty, two morphological features that have been associated with both elements. 2. We use stable carbon isotope ratios of faeces to reconstruct diet in terms of proportions of C(3) plants (browse) and C(4) plants (grass) consumed by different species in different seasons. These data are combined with proxies for diet quality (per cent nitrogen, neutral detergent fibre, acid detergent fibre, and acid detergent lignin) from faeces to track changes in diet quality. 3. Two statistical approaches are used in model selection, i.e. tests of significant correlations based on linear regression analyses, and an information-theory approach (Akaike's Information Criterion) providing insight into strength of evidence for models. 4. Results of both methods show that, contrary to many predictions, body mass and diet type are not related, but these data confirm predictions that diet quality decreases with increasing body size, especially during the dry season. Hypsodonty, as expected, varies with diet type, increasing with increased grass intake. 5. These findings support both a diet type and diet quality model, implying some degree of exclusivity. We propose that congruence between models may be achieved through addition of diet quality proxies not included here, because hypsodonty is more likely a reflection of the abrasive properties of consumed foods, i.e. related to food quality, rather than food type. This implies that adaptation to diets of varying quality, through changes in body size and dental features, has been the primary mechanism for diversification in ungulates. 6. Our interpretation contrasts with several recent studies advocating diet type as the primary factor, exemplifying that further reconciliation between the two models is needed. We discuss the implications of this study for future approaches to achieve a more cohesive understanding of the evolutionary outcomes of herbivore nutrition.     

Effect of dietary lipid content on lipid transport and storage during larval development of Manduca sexta

Manduca sexta larvae were raised on diets containing either 1.2% fat (control diet), 5.9% fat (high-fat diet) or on a fat-free diet. Insects raised on the control and high-fat diets did not differ significantly in body weight, whereas animals raised on the fat-free diet were significantly smaller. The fat content of the diet had no effect on the hemolymph concentration of lipophorin. During the larval period, lipophorin isolated from animals on the high-fat diet contained more lipid, and lipophorin isolated from animals on the fat-free diet contained less lipid than lipophorin isolated from control animals. However, lipophorin isolated from animals during the prepupal period had the same composition regardless of diet. Compared to controls, animals on the high-fat diet had a larger mass of fat body which contained more stored triacylglycerol, while animals on the fat-free diet had a smaller mass of fat body which contained less stored triacylglycerol. As the fat content of the diet was increased, the fatty acid composition of fat body triacylglycerols reflected more closely that of the dietary lipid.     

Which body condition index is best?

Body condition indices are widely used by ecologists, but many indices are used without empirical validation. To test the validity of a variety of indices, we compared how well a broad range of body condition indices predicted body fat mass, percent body fat and residual fat mass in mice Mus musculus. We also compared the performance of these condition indices with the multiple regression of several morphometric variables on body fat mass, percent body fat and residual fat mass. In our study population, two ratio based condition indices – body mass/body length and log body mass/log body length – predicted body fat mass as well as or better than other ratio and residual indices of condition in females. In males one ratio based condition index (log body mass/log body length) and one residual index (residuals from a regression of pelvic circumference on body length) were best at predicting body fat mass. All indices were better at estimating body fat mass, and residual fat mass than at estimating percent body fat. The predictions of body fat were much better for females than for males. Multiple regressions incorporating pelvic circumference (i.e. girth at the iliac crests) were the best predictors of body fat mass, residual fat mass, and percent body fat, and these multiple regressions were better than any of the condition indices. We recommend 1) that condition be precisely defined, 2) that predictors of condition be empirically validated, 3) that pelvic circumference be considered as a potential predictor of fat content, and 4) that, in general, multiple regression be considered as an alternative to condition indices.     

APPRAISAL OF CONDITION ESTIMATION METHODS FOR EAST AFRICAN UNGULATES

Previously proposed methods of determining physical condition of game animals are discussed. A total of 286 African ungulates of ten species were collected and autopsied, and several indices of physical condition were calculated. Selected indices were correlated with total fat or total lean percentages or both. Of the methods discussed and evaluated, those based upon fat deposited in particular areas were more accurate or feasible than those based upon blood constituents, adrenal weights, or body weight/skeletal measurement ratios.     

Effect of diet on condition indices in black-tailed jackrabbits

Changes in blood, urine and physical condition indices in 23 adult male black-tailed jackrabbits (Lepus californicus) with ad libitum feeding and 25% feed restriction were measured over a 2 wk period from 30 May to 12 June 1988. Feed restricted jackrabbits had (1) lower post-trial body weights and kidney fat indices, (2) higher femur marrow fat, serum bilirubin and cortisol concentrations, and adrenal cortex width, and (3) depressed immune function. No single index alone could best measure the nutritional status of these jackrabbits.     

Influence of two different diets on fluid and particle retention time Javan langur (Trachypithecus auratus auratus)

In captivity, langurs (foregut fermenting primates) often suffer from digestive disorders, and in particular display soft stools on diets with a high proportion of fruits, vegetables, and grain products. In this study, we tested whether the improvement in feces consistency expected after an omission of vegetables from a conventional diet was also accompanied by a change in the excretion pattern as measured by the mean retention time (MRT). Two adult Javan langurs were kept together and offered a conventional diet of pellets, browse, vegetables, or a diet consisting of pellets and browse only. MRT were measured with cobalt-EDTA as the fluid and chromium oxide as the particle marker before, during, and after the test diet, collecting the feces of both animals together. MRTs of fluids and particles were longer (47/49 h) on the test diet as compared to the usual diet (42/43 h). Feces consistency improved during the test period. The test period was marked by an increased fiber intake; however, on the conventional diet, dietary water intake exceeded the calculated water flux for these animals distinctively, due to the high proportion of vegetables. Therefore, it cannot be decided whether the increase of fiber level alone or also a reduction of the excessive water intake was responsible for the changes observed. Comparing the results of this study to other studies on ingesta retention in foregut fermenting primates, no correlation between the body mass of the animals and the MRTs measured is evident, which is in accordance with observations in other groups of foregut fermenters.     

The effect of diet quality on physiological and life-history traits in the harvestman Pachylus paessleri

The balance between energy acquisition and expenditure is critical to the survival and reproductive success of animals. Here we investigate the long-term effects of diet quality on physiological and life-history flexibility in the harvestman, Pachylus paessleri. We used cow meal as a protein-rich diet and potatoes as a carbohydrate-rich diet in order to reproduce two extreme conditions regarding food quality in harvestmen natural habitat. As proxy variables of the energy expenditure process, we quantified standard metabolic rate (maintenance), changes in body mass (somatic condition), and fecundity (reproduction). We found that animals consuming the protein-rich diet were able to increase both their body condition and fecundity. However, the increment in these two life-history traits was correlated with higher maintenance costs. In contrast, the carbohydrate-rich diet did not provide enough specific nutrients for reproductive events, although it may have allowed animals to survive for a long time. Thus, according to the quality of the diet available in the environment, harvestman females can adopt different life-history strategies correlated with phenotypic adjustments at anatomical and physiological levels. In the Mediterranean region, spatial and temporal changes in food quality are typical, so greater phenotypic flexibility is expected to cope with this kind of environmental variation.     

Impact of nutrition on oocyte quality: cumulative effects of body composition and diet leading to hyperinsulinemia in cattle

The present study sought to assess the combined effects of body composition and diet (level of feeding) on the postfertilization developmental potential of oocytes recovered from heifers using ultrasound-guided transvaginal follicular aspiration and to relate oocyte quality to the metabolic status of these animals. By collecting oocytes on repeated occasions spanning several weeks, it was possible to assess the cumulative effects of changes in nutritional status on oocyte quality over this period. Twenty-four heifers of low and moderate body condition were placed on one of two levels of feeding (equivalent to once or twice the maintenance requirements of these animals). Oocytes were recovered at two defined time points within each of three successive estrous cycles and were matured, fertilized, and cultured to the blastocyst stage in vitro. The results show that the effect of feeding level on oocyte quality is dependent on the body condition of the animal, with the high level of feeding being beneficial to oocytes from animals of low body condition but detrimental to oocytes from animals of moderately high body condition. Furthermore, the effects of high levels of feeding on oocyte quality were cumulative, with blastocyst yields for relatively fat heifers on twice the maintenance requirement deteriorating with time relative to yields for relatively thin heifers on the same level of feeding. Finally, a significant proportion of the moderately fat animals on the high level of feeding were hyperinsulinemic, and we show, to our knowledge for the first time in ruminants, that this condition is associated with impaired oocyte quality.     

光周期和高脂食物对雌性高山姬鼠能量代谢和产热的影响

为了研究光周期和高脂食物对小型哺乳动物能量代谢和产热的影响,将成年雌性高山姬鼠分别驯化于长光照低脂、高脂食物和短光照低脂、高脂食物条件下,7周后测定动物的体重、能量摄入、产热、身体组成、血清瘦素浓度以及体脂含量等参数。结果发现:1)短光照抑制体重增长,降低血清瘦素浓度,增加非颤抖性产热;2)高脂食物使摄入能减少,消化率和体脂含量提高,但未显著影响体重、基础代谢率、非颤抖性产热和血清瘦素浓度;3)血清瘦素浓度与摄入能不相关,但与体脂重量正相关。结果暗示:短光照下瘦素作用敏感性增加和产热能力增强,可能介导了抵抗高脂食物诱导的肥胖。在野外条件下,高山姬鼠能通过能量代谢和产热的适应性调节避免体重的过度增长,有利于降低捕食风险,增强生存能力。     

Effects of grass and browse consumption on the winter mass dynamics of elk

Much of the research into herbivore ecology and evolution has focused on patterns and mechanisms of niche partitioning in the diets of specialized grazers and browsers, but the significance of the balance of grazing and browsing within intermediate feeders present unique questions that have received less attention. We explored the nutritional effects of varying the balance of grass and browse in the winter diets of elk (Cervus elaphus). We compiled published data from three similar studies that monitored the mass dynamics of captive elk fed diets of pure grass, pure browse or 1 of 14 mixed diets in winter feeding trials. Elk lost mass (up to 22% of initial body weight) in 29 of 33 feeding trials, similar to wild elk in winter. We used regression models of mass dynamics, considering the linear, quadratic or logarithmic effects of the proportion of the diet that was grass (≈1 − proportion of the diet that was browse) and the additive and interactive effects of nitrogen intake. Diet composition had strong effects on mass dynamics, and all models explained ≥73% of the variation (adjusted r ²) in mass dynamics. Nitrogen intake had uniformly positive effects on mass balance, and increasing grass intake caused improved maintenance of body mass up to a point, but further increases in the proportion of grass in the diet had neutral or negative effects on body mass. Overall, the data suggest that elk are adapted to consuming mixed diets. Nonetheless, data on the foraging behavior and diet selection of wild elk in many populations show that elk often consume grass-dominated diets and sometimes consume browse-dominated diets, but rarely consume mixed diets. Physiological adaptations to mixed diets may place unique spatio-temporal constraints on diet selection in intermediate feeders and impose large penalties for a sub-optimal balance of grazing and browsing.     

Herbivore Body Condition Response in Altered Environments: Mule Deer and Habitat Management

The relationships between habitat, body condition, life history characteristics, and fitness components of ungulates are interwoven and of interest to researchers as they strive to understand the impacts of a changing environment. With the increased availability of portable ultrasound machines and the refinement of hormonal assays, assessment of ungulate body condition has become an accessible monitoring strategy. We employed body condition scoring, estimation of % ingesta-free body fat (%IFBF), assessment of free thyroid hormones (FT4 and FT3), and assessment of pregnancy, as metrics to determine if landscape-level habitat treatments affected body condition of adult (≥1.5 years old) female mule deer (Odocoileus hemionus). All body condition related metrics were measured on 2 neighboring study areas — a reference area that had received no habitat treatments and a treatment study area that had received mechanical removal of pinyon pine (Pinyus edulis) - Utah juniper (Juniperus osteosperma) forest, chemical control of weeds, and reseeding with preferred mule deer browse species. A consistent trend of higher %IFBF was observed in the treatment study area than in the reference study area , although variation of estimates was larger than hypothesized. A similar pattern was observed with higher thyroid hormones concentrations being observed in the treatment study area, but large amounts of variation within concentration estimates were also observed. The consistent pattern of higher body condition related estimates in our treatment study area provides evidence that large mammalian species are sensitive to landscape change, although variation within estimates underlie the challenge in detecting population level impacts stemming from environmental change.     

Landscape fragmentation influences winter body mass of roe deer

Body size of large herbivores is a crucial life history variable influencing individual fitness‐related traits. While the importance of this parameter in determining temporal trends in population dynamics is well established, much less information is available on spatial variation in body size at a local infra‐population scale. The relatively recent increase in landscape fragmentation over the last century has lead to substantial spatial heterogeneity in habitat quality across much of the modern agricultural landscape. In this paper, we analyse variation in body mass and size of roe deer inhabiting a heterogeneous agricultural landscape characterised by a variable degree of woodland fragmentation. We predicted that body mass should vary in relation to the degree of access to cultivated meadows and crops providing high quality diet supplements. In support of our prediction, roe deer body mass increased along a gradient of habitat fragmentation, with the heaviest deer occurring in the most open sectors and the lightest in the strict forest environment. These spatial differences were particularly pronounced for juveniles, reaching >3 kg (ca 20% of total body mass) between the two extremes of this gradient, and likely have a marked impact on individual fates. We also found that levels of both nitrogen and phosphorous were higher in deer faecal samples in the more open sectors compared to the forest environment, suggesting that the spatial patterns in body mass could be linked to the availability of high quality feeding habitat provided by the cultivated agricultural plain. Finally, we found that adults in the forest sector were ca 1 kg lighter for a given body size than their counterparts in the more open sectors, suggesting that access to nutrient rich foods allowed deer to accumulate substantial fat reserves, which is unusual for roe deer, with likely knock‐on effects for demographic traits and, hence, population dynamics.     

Seasonal fluctuations of body condition in northern sika deer on Hokkaido Island, Japan

We analyzed seasonal and sexual fluctuations in kidney mass (KM) and kidney fat mass (KFM) as indices of condition in Hokkaido sika deerCervus nippon yesoensis Heude, 1884. For 76 male and 132 female sika deer, seasonal fluctuations in KM and KFM were given by fitted sine wave growth curves. Although the kidney fat index (KFI) is used frequently to evaluate animal condition, we reject it because it is based on the assumption that kidney mass is proportional to body mass in all seasons. Our data did not support this assumption. KFM is a better indicator of Hokkaido sika deer condition than KFI. Although sex-based differences in cervid KFM are said to reflect differences in reproductive cycles, the seasonal similarities in sika deer KFM levels may represent adaptations to the long severe Hokkaido winter. Because in our study deer populations were at low densities and had high pregnancy rates, our sine wave growth models can be regarded as reference for fat level fluctuations in Hokkaido sika deer.     

Matrix Intensification Affects Body and Physiological Condition of Tropical Forest-Dependent Passerines

Matrix land-use intensification is a relatively recent and novel landscape change that can have important influences on the biota within adjacent habitat patches. While there are immediate local changes that it brings about, the influences on individual animals occupying adjacent habitats may be less evident initially. High-intensity land use could induce chronic stress in individuals in nearby remnants, leading ultimately to population declines. We investigated how physiological indicators and body condition measures of tropical forest-dependent birds differ between forest adjacent to surface mining sites and that near farmlands at two distances from remnant edge in southwest Ghana. We used mixed effects models of several condition indices including residual body mass and heterophil to lymphocyte (H/L) ratios (an indicator of elevated chronic stress) to explore the effect of matrix intensity on forest-dependent passerines classed as either sedentary area-sensitive habitat specialists or nomadic generalists. Individual birds occupying tropical forest remnants near surface mining sites were in poorer condition, as indicated by lower residual body mass and elevated chronic stress, compared to those in remnants near agricultural lands. The condition of the sedentary forest habitat specialists white-tailed alethe, Alethe diademata and western olive sunbird, Cyanomitra obscura was most negatively affected by high-intensity surface mining land-use adjacent to remnants, whereas generalist species were not affected. Land use intensification may set in train a new trajectory of faunal relaxation beyond that expected based on habitat loss alone. Patterns of individual condition may be useful in identifying habitats where species population declines may occur before faunal relaxation has concluded.