Interactions among host diet, nutritional status and gastrointestinal parasite infection in wild bovids

In this study, I explored the interactions among host diet, nutritional status and gastrointestinal parasitism in wild bovids by examining temporal patterns of nematode faecal egg shedding in species with different diet types during a drought and non-drought year. Study species included three grass and roughage feeders (buffalo, hartebeest, waterbuck), four mixed or intermediate feeders (eland, Grant's gazelle, impala, Thomson's gazelle) and two concentrate selectors (dik-dik, klipspringer). Six out of the nine focal species had higher mean faecal egg counts in the drought year compared to the normal year, and over the course of the dry year, monthly faecal egg counts were correlated with drought intensity in four species with low-quality diets, but no such relationship was found for species with high-quality diets. Comparisons of dietary crude protein and faecal egg count in impala showed that during the dry season, individuals with high faecal egg counts (> or =1550 eggs/g of faeces) had significantly lower crude protein levels than individuals with low (0-500 eggs/g) or moderate (550-1500 eggs/g) egg counts. These results suggest that under drought conditions, species unable to maintain adequate nutrition, mainly low-quality feeders, are less able to cope with gastrointestinal parasite infections. In particular, during dry periods, reduced protein intake seems to be associated with declining resilience and resistance to infection.     

Effects of dehydration and heat stress on food intake and dry matter digestibility in East African ruminants

Comparative investigations were made between wild and domestic ruminants from arid and semi-arid regions and those species from non-arid areas in an attempt to evaluate the adaptations of these ruminants in terms of the effects of heat stress and dehydration on food intake and digestibility. The effect of (a) an intermittent heat load (a daily light cycle of 12 h at 22 degrees C and 12 h at 40 degrees C) compared to 22 degrees C throughout the day and (b) dehydration level of 15% weight loss, with and without the heat load, on the intake and digestibility of a poor quality hay was investigated in the Grant's gazelle, Oryx, the domestic Turkana goats, fat-tailed sheep, zebu cattle, Thomson's gazelle and wildebeest. The intermittent heat load with water available ad libitum depressed the food intake of zebu cattle and Turkana goats by more than 40%. It had no significant effect on the food intake of the other species. The Thomson's and Grants gazelle, oryx, wildebeest and fat-tailed sheep appear well adapted to withstanding a periodic heat load. Dehydration at 22 degrees C caused a marked depression on food intake of all the species investigated. Dehydration together with a heat load caused no further reduction in the food intake by the Grants's gazelle, oryx, and goats but it did cause a further reduction in the intake in the other species. The small non-domestic ruminants (i.e. Grant's and Thomson's gazelle) appear much more digestive efficient than any of their domestic counterpart.     

Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system

Summary A review is made of the ruminant digestive system in its morphophysiological variations and adaptations relating to foraging behaviour, digestive physiology, to interactions between plants and ruminants and to geographic and climatic diversity of ruminants' ecological niches. Evidence is provided for evolutionary trends from an extreme selectivity mainly for plant cell contents and dependence upon a fractionated fore- and hindgut fermentation, to an unselective intake of bulk roughage subjected to an efficient plant cell wall fermentation, mainly in the forestomachs. The review is based on detailed comparative morphological studies of all portions of the digestive system of 65 ruminant species from four continents. Their results are related to physiological evidence and to the classification of all extant ruminants into a flexible system of three overlapping morphophysiological feeding types: concentrate selectors (40%), grass and roughage eaters (25%) and intermediate, opportunistic, mixed feeders (35%). Several examples are discussed how ruminants of different feeding types are gaining ecological advantage and it is concluded that ruminants have achieved high levels of digestive efficiency at each evolutionary stage, (including well-documented seasonal adaptations of the digestive system) and that ruminant evolution is still going on. Deductions made from the few domesticated ruminant species may have, in the past, biased scientific evaluation of the free-ranging species' ecology. The main threat to a continuous ruminant evolution and diversity appears to be man's neglect for essential ecological interactions between wild ruminants and their specific habitats, which he alters or destroys.Abbreviations bw body weight - CS concentrate selector - DFC distal fermentation chamber (distended caecocolon) - GR grass and roughage eater - IM intermediate (mixed) feeder - PFC proximal fermentation chamber (ruminoreticulum/forestomachs) - RR Ruminoreticulum - SCFA Short-chain fatty acis (acetic, butyric, propionic acid set free by rumen bacteriae) - SE Surface enlargement (of absorptive mucosa) Supported by German Research Community grant DFG Ho 273/6Dedicated to Professor Dr. Dr. h.c. Dietrich Starck on the occasion of his 80th birthday     

Ruminant diversification as an adaptation to the physicomechanical characteristics of forage.

The validity of Hofmann's classification of ruminants into browsers/"concentrate selectors", intermediate feeders and grazers/"grass and roughage eaters" and of his consecutive physiological postulates has repeatedly been questioned. In contrast to former concepts, which all focused on the chemical characteristics of the respective forages, we propose a new hypothesis on the main driving force of ruminant diversification, namely the physicomechanical characteristics of the respective forages. In contrast to browse, grass tends to stratify and form a "fibrous raft" in the reticulorumen. The significantly more capacious forestomachs of grazers, and the significantly thicker rumen pillars (indicating the strength of reticulorumen muscle equipment) of their forestomachs, are interpreted as particular adaptations to this forage characteristic. With these parameters, we present, for the first time, two single morphological measurements that allow the statistical reconstruction of Hofmann's classification. A small forestomach capacity and the lack of strong reticulorumen muscles in browsers would explain the observed exclusiveness with which browsers avoid grass forage under natural conditions, which we confirmed using two datasets on the composition of the natural diet. Both rumen pillar thickness and relative forestomach capacity were significantly correlated to the grass content of the natural diet, respectively. Our functional interpretation was also supported by a stepwise regression analysis with the proportion of grass in the natural diet as dependent variable and the rumen pillar thickness, the relative forestomach capacity, and the body weight as independent variables, which revealed significant equations.     

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.     

The biomass of game animals in Nairobi National Park, 1960–66

The large animals of Nairobi National Park have been censused intermittently once a month from mid-1960 until the end of 1966. Of the 23 species discussed, the density per kilometre has been determined accurately for the following: Grant's gazelle, Thomson's gazelle, water-buck, impala, wildebeest, hartebeest, eland, buffalo, warthog, giraffe, zebra and ostrich. From these figures the biomass has been calculated for each species each year. After a peak biomass of 12,775 kg/sq. km during the drought of 1960–61 the population declined to and remained fairly level at about 5690 kg/sq. km from 1962 to 1966. This figure includes about 1050 kg/sq. km of cattle and sheep. The large predators account for only about 1.4 % of the total biomass of wild animals while removing about 15.5% from this group annually.     

The relative roles of body size and feeding type on activity time of temperate ruminants

Recently, there has been some critical testing of whether body size or feeding type (CS: concentrate selectors, IF: intermediate feeders, GR: grass-roughage eaters) is the most important determinant of physiological aspects of ruminant ecology, whereas little has been done on behavioral aspects like activity time. Different predictions regarding the relationships between activity time and body weight/feeding type were tested with activity time data from 18 temperate ruminants. Activity time decreased allometrically with increasing body weight, but there was also a tendency for an effect of feeding type. Exclusion of one statistically defined outlier (mountain goat) made the effect of feeding type highly significant. GR and CS were about equally active. Surprisingly, IF were more active than both GR and CS. The hypothesis is put forward that IF are more active than GR/CS due to their opportunistic use of high-quality forage of both types (concentrate and grass-roughage; on average better quality and hence shorter rumination time), though possible confounding effects of observation methods and varied behavior with respect to cover among CS, IF, and GR should also be evaluated. Received: 22 May 1997 / Accepted: 22 September 1997     

Anatomy of the tongue and microstructure of the lingual papillae in the fallow deer Dama dama (Linnaeus, 1758)

The function of mouth organs in ruminants is connected with the process of rumination. To study morphofunctional relations, microstructures in tongues of 4- to 5-year old adult fallow deer were examined using scanning electron microscopy.When analyzing the tongue of the fallow deer, i.e. a ruminant classified as an intermediate mixed feeder between grass and roughage eaters, two processes were taken into account: (i) foraging and forage selecting, and (ii) chewing the cud during rumination to reduce particle size and improve digestibility.Microstructural results show that the above mentioned processes in fallow deer are important selection factors, which in the anterior part of tongue led to the development of clusters of fungiform papillae connected with preselection of food as well as a specific pattern of filiform papillae promoting increased adhesion of transported food. Massive and flattened conical papillae on the torus are arranged according to sideways jaw movements and are co-localized with flattened fungiform papillae and two rows of vallate papillae. Such an arrangement of papillae on the lingual torus presumably facilities distribution of ruminated food, with simultaneous transferring of taste signals about masticated food particles.     

Food preferences and tooth wear in the sand gazelle (Gazella marica)

Food preferences of the sand gazelle (Gazella marica) from the Mahazat as-Sayd Protected Area in Saudi Arabia were evaluated using focal animal sampling in conjunction with an eco-morphological method examining two parameters of tooth wear, i.e., occlusal relief and cusp shape. Observations of live, free-ranging animals (n = 53) showed that sand gazelles generally consumed more grass (58.4%) than browse (41.6%). However, during the dry season, gazelles spent significantly more time browsing (51.0%) and less time grazing (49.0%) than under wet conditions (browsing: 17.6%; grazing: 82.4%). Thus, consistent with predictions, sand gazelles are intermediate feeders but shift towards browsing when grass is scarce. The mesowear signature of the sand gazelle is consistent with a grazing signal in other ruminants. In other words, the browse component of the diets of live animals was not reflected in the tooth wear. This could have occurred because browse is less abrasive than grass, but more likely because all food types are heavily abrasive in this dusty habitat. We conclude that the sand gazelle population in Mahazat as-Sayd encounters a highly abrasive diet, which has implications for their ability to meet nutritional demands.     

Activity of cellulolytic enzymes in the contents of reticulorumen and caecocolon of roe deer (Capreolus capreolus)

Selective ruminants, which prefer easily digestible plants, cannot digest fibrous forage as well as grass eaters. Low enzyme activity or short retention time of ingesta particles in fermentation chambers appeared to be responsible for reduced cellulose breakdown. Seasonal activity of cellulolytic enzymes, cellulose concentration and protozoa population in reticulorumen (RR) and caecocolon (CC) of roe deer as a typical concentrate selector were investigated. Cellulase activities were lowest in winter when cellulose concentration in RR contents were highest. Highest enzyme activities and lowest cellulose concentration were measured in early spring. Cellulolytic activities were significantly correlated with the number of protozoa in RR. Only one entodinomorphic genus was identified in the RR. The enzyme activities in CC were far lower compared with those in RR. Low cellulose digestion in the RR cannot be compensated for by cellulose breakdown in the CC. The reduced cellulose digestion of roe deer may be attributed to the short retention time of food particles in spring and summer, whereas decreased colonisation of microorganisms in the rumen may be the main reason for low cellulose breakdown in winter.     

Rumen Ciliate Protozoa of the Blue Duiker (Cephalophus monticola), with Observations on Morphological Variation Lines Within the Species Entodinium dubardi

ABSTRACT. Protozoal concentrations were determined in rumen and cecal contents of 20 blue duikers ( Cephalophus monticola ). Ten animals of each sex were fed either a high concentrate or high roughage diet. Rumen protozoa were present in 19 of the 20 animals and concentrations ranged from 4.5 to 33.7 × 10⁶ per g of rumen contents. At the higher concentrations, protozoal cells equaled between 30–40% of the total rumen contents volume. No protozoa were found in cecal contents. Weight of rumen contents was higher in females than in males ( P < 0.01), and rumen protozoa concentrations were higher in males ( P < 0.05) and in those animals fed the high concentrate diet ( P < 0.05). All the protozoa were identified as belonging to a single species, Entodinium dubardi . However, an average of about 30% of the E. dubardi cells varied from the typical morphology of this species. These cells appeared to be on variation lines leading toward 7–10 other non-caudate species of Entodinium . The present data were used to evaluate and discuss the concept of variation lines within E. dubardi .     

Use of δ13C values to determine vegetation selectivity in East African herbivores

Summary The quantitative plant species composition of the rumen contents of a large number of individuals from eight East African herbivores was determined by direct visual analysis. All plant species were classified as either C₃ or C₄, and an estimated ₁₃C for the rumen sample was calculated. This estimated value was compared to a measured value determined directly from rumen subsample. The two methods of determining quantitative C₃ and C₄ composition differed by less than 1%, and the isotopic analysis has the advantage of being rapid and totally objective.The isotopic analysis allowed us to differentiate between grazers and browsers and to determine the quantitative dependence of each animal on C₃ and C₄ photosynthetic types. Kongoni, wildebeest, cattle, and sheep were nearly pure grazers on the Athi Kapiti Plains; and the Grant's gazelle were predominantly browsers. Thompson's gazelle, goast and impala were intermediate. The species most dependent upon browse showed a marked and rapid shift to grass within a few days following rain. This isotopic method may have general utility in the study of East African ecology.     

Evaluation of the effects of dietary particle fractions on fermentation profile and concentration of microbiota in the rumen of dairy cows fed grass silage-based diets

The study evaluated the effects of three different theoretical particle lengths (TPL) of grass silage on the distribution of particle fractions of the diet and the resulting effects on fermentation profile and concentrations of protozoa and mixed bacterial mass in the rumen of three lactating Holstein cows fed total mixed rations (45% grass silage, 5% grass hay and 50% concentrate) ad libitum. Decreasing TPL of grass silage (long, medium, short) reduced particles retained on the 19-mm sieve of the Penn State Particle Separator, while particle fractions from 8 mm to 19 mm and smaller than 8 mm were increased. Different TPL did not affect pH and the concentration of volatile fatty acids in the rumen. However, lowering the TPL from long to medium increased significantly the bicarbonate concentration, acetate proportion and protozoal number in the rumen, whereas the proportion of bacterial protein in ruminal digesta and its amino acid concentration were significantly increased by the short TPL. For the current feeding conditions, it can be concluded that increasing the fraction of particles between 8 and 19 mm and probably even the fraction below 8 mm by decreasing TPL of grass silage do not adversely affect rumen conditions and can be beneficial in terms of optimising concentration and activity of ruminal microbiota in high-yielding dairy cows.     

Evaluation of the effects of dietary particle fractions on fermentation profile and concentration of microbiota in the rumen of dairy cows fed grass silage-based diets

The study evaluated the effects of three different theoretical particle lengths (TPL) of grass silage on the distribution of particle fractions of the diet and the resulting effects on fermentation profile and concentrations of protozoa and mixed bacterial mass in the rumen of three lactating Holstein cows fed total mixed rations (45% grass silage, 5% grass hay and 50% concentrate) ad libitum. Decreasing TPL of grass silage (long, medium, short) reduced particles retained on the 19-mm sieve of the Penn State Particle Separator, while particle fractions from 8 mm to 19 mm and smaller than 8 mm were increased. Different TPL did not affect pH and the concentration of volatile fatty acids in the rumen. However, lowering the TPL from long to medium increased significantly the bicarbonate concentration, acetate proportion and protozoal number in the rumen, whereas the proportion of bacterial protein in ruminal digesta and its amino acid concentration were significantly increased by the short TPL. For the current feeding conditions, it can be concluded that increasing the fraction of particles between 8 and 19 mm and probably even the fraction below 8 mm by decreasing TPL of grass silage do not adversely affect rumen conditions and can be beneficial in terms of optimising concentration and activity of ruminal microbiota in high-yielding dairy cows.     

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.     

Ehrlichia ruminantium infection (heartwater) in wild animals

Several wild animal species have been implicated as hosts of Ehrlichia ruminantium (formerly Cowdria ruminantium), the rickettsial agent causing heartwater, a fatal disease of domestic ruminants in sub-Saharan Africa and eastern Caribbean. However, evidence for infection in most wild species is inconclusive because of inadequate diagnostic techniques. Infection has been proven only in 12 African ruminants, three non-African ruminants and two African rodents. A subclinical carrier state occurs in eight of the African ruminant species. Further studies on E. ruminantium infection in wild animal species are needed in order to determine the host range of this pathogen accurately. The host range of Ehrlichia ruminantium in wildlife is reviewed here and the role played by these species in the epidemiology and spread of heartwater is discussed.     

Influence of ciliate protozoa on biochemical changes and hydrolytic enzyme profile in the rumen ecosystem

AIMS: To assess the effect of presence or absence of rumen protozoa on fermentation characteristics and enzyme profile in growing lambs. METHODS AND RESULTS: Weaner lambs (G1, G2, G3, G4, G5 and G6 groups) were defaunated by oral administration of sodium laurel sulphate (at 8 g 100 kg(-1) body weight). The lambs of G4, G5 and G6 groups were refaunated. The roughage and concentrate ratio in the diet of G1 and G4, G2 and G5, and G3 and G6 were 50:50 (R1), 65:35 (R2) and 80:20 (R3), respectively. Daily dry matter intake was similar in defaunated and faunated lambs. However, digestibility of organic matter (OM), cellulose and gross energy were lower in defaunated lambs while crude protein (CP) digestibility was similar in both defaunated and faunated lambs. The rumen pH and NH3-N were lower (P < 0.01) while TVFA, total-N and TCA-ppt-N were higher (P < 0.01), in defaunated lambs. Ruminal activity of carboxymethyl cellulase was lower (P < 0.01) in defaunated lambs and amylase, xylanase, protease and urease were similar in faunated and defaunated lambs. Nutrient utilization, rumen metabolites and ciliate protozoal count were higher, whereas digestibility of fibre fractions was lower in high rather than low concentrate fed lambs. The rumen protozoa present before defaunation were B-type and the protozoa which re-established on refaunation were also B-type. CONCLUSIONS: Absence of ciliate protozoa decreased nutrient digestibility and increased ruminal TVFA and total-N with lower NH3-N concentration, indicating better energy and protein utilization in defaunated lambs. SIGNIFICANCE AND IMPACT OF THE STUDY: Defaunation improved energy and protein utilization in lambs.     

Molar proportions of volatile fatty acids in the gastrointestinal tract of East African wild ruminants

The molar proportions of seven individual VFA's were determined at select sites along the gastrointestinal tract of sixteen species of East African wild ruminants. The resulting data were statistically analyzed for species effect, and for effects due to major feeding groups (browsers, grazers, fresh grass grazers, etc.) and for body weight groups (5-750 kg animals). Present data suggest that body weight, rather than diet, is the more influential factor in reticulo-rumen fermentation rate, and in the molar proportion of fatty acids present. The molar proportions of VFA's observed in the mid and hindgut of these wild ruminants appeared more responsive to diet and body weight of the animal than did foregut VFA values.     

Culicoides Species Communities Associated with Wild Ruminant Ecosystems in Spain: Tracking the Way to Determine Potential Bridge Vectors for Arboviruses

The genus Culicoides Latreille 1809 is a well-known vector for protozoa, filarial worms and, above all, numerous viruses. The Bluetongue virus (BTV) and the recently emerged Schmallenberg virus (SBV) are responsible for important infectious, non-contagious, insect-borne viral diseases found in domestic ruminants and transmitted by Culicoides spp. Both of these diseases have been detected in wild ruminants, but their role as reservoirs during the vector-free season still remains relatively unknown. In fact, we tend to ignore the possibility of wild ruminants acting as a source of disease (BTV, SBV) and permitting its reintroduction to domestic ruminants during the following vector season. In this context, a knowledge of the composition of the Culicoides species communities that inhabit areas where there are wild ruminants is of major importance as the presence of a vector species is a prerequisite for disease transmission. In this study, samplings were conducted in areas inhabited by different wild ruminant species; samples were taken in both 2009 and 2010, on a monthly basis, during the peak season for midge activity (in summer and autumn). A total of 102,693 specimens of 40 different species of the genus Culicoides were trapped; these included major BTV and SBV vector species. The most abundant vector species were C. imicola and species of the Obsoletus group, which represented 15% and 11% of total numbers of specimens, respectively. At the local scale, the presence of major BTV and SBV vector species in areas with wild ruminants coincided with that of the nearest sentinel farms included in the Spanish Bluetongue Entomological Surveillance Programme, although their relative abundance varied. The data suggest that such species do not exhibit strong host specificity towards either domestic or wild ruminants and that they could consequently play a prominent role as bridge vectors for different pathogens between both types of ruminants. This finding would support the hypothesis that wild ruminants could act as reservoirs for such pathogens, and subsequently be involved in the reintroduction of disease to livestock on neighbouring farms.