Dietary niche, feeding type classification and seasonal strategies of Rangifer tarandus and Ovibos moschatus discussed in relation to the anatomy of their digestive system. Classification criteria for the flexible feeding type system, originally established in bovids and later adapted to cervids, are substantiated and critically discussed in the light of recent attempts to invalidate the system. Eurasian mountain reindeer, North American barren-ground caribou, Svalbard reindeer and Victoria Island caribou are seasonally adaptable, opportunistic ruminants of the intermediate feeding type but the long evolutionary sepatation of Svalbard reindeer has modified several morphological features for winter survival without lichen, resulting in better adaptation to fibrous forage. Muskoxen, despite their seasonal selectivity, are typical grass and roughage eaters with extremely long mean retention time. Detailed data on the entire digestive system from muzzle to anus on both species are still insufficient and extended studies are worthwhile for understanding their nutritional niche and feeding adaptations.

The phylogenetic relationships of Hunter's antelope or hirola Beatragus hunteri (Artiodactyla: Alcelaphini), one of the most seriously threatened antelopes in Africa, have been the subject of controversial discussion. A total of 801 base pairs of the mitochondrial cytochrome b gene were determined from five alcelaphine species and from two nonalcelaphine bovids. Phylogenetic analysis using parsimony, maximum likelihood, and distance methods identified a single best-supported hypothesis of evolutionary relationships within the Alcelaphini: (i) monophyly of the tribe, (ii) most basal position of the blue wildebeest, (iii) sister-grouping of the Lichtenstein's and Coke's hartebeests, (iv) phylogenetic distinctiveness of the hirola from the topi and the hartebeests. We deduce that the alcelaphine cytochrome b sequences began to diverge from a common mitochondrial ancestor about 5.5 million years ago. In addition, we provide molecular evidence that the impala Aepyceros melampus is not closely related to Alcelaphini.

The reticulum is the second part of the ruminant forestomach, located between the rumen and the omasum and characterized by honeycomb-like internal mucosa. With its fluid contents, it plays a decisive role in particle separation. Differences among species have been linked to their feeding style. We investigated whether reticulum size (absolute and in relation to rumen size) and size of the crests that form the mucosal honeycomb pattern differ among over 60 ruminant species of various body sizes and feeding type, controlling for phylogeny. Linear dimensions generally scaled allometrically, that is to body mass0.33. With or without controlling for phylogeny, species that ingest a higher proportion of grass in their natural diet had both significantly larger (higher) rumens and higher reticular mucosa crests, but neither reticulum height nor reticulum width varied with feeding type. The height of the reticular mucosa crests represents a dietary adaptation in ruminants. We suggest that the reticular honeycomb structures do not separate particles by acting as traps (neither for small nor for large particles), but that the structures reduce the lumen of the reticulum during contractions - at varying degrees of completeness in the different feeding types. In browsing species with rumen contents that may be less fluid and more viscous than those of the reticulum, incomplete closure of the lumen may allow the reticulum to retain the fluid necessary for particle separation. In grazing species, whose rumen contents are more stratified with a larger distinct fluid pool, a more complete closure of the reticular lumen due to higher crests may be beneficial as the reticulum can quickly re-fill with fluid rumen contents that contain pre-sorted particles.

Using cranioskeletal measurements, several studies have generated evidence that grazing ruminants have a more pronounced mastication apparatus, in terms of muscle insertion areas and protuberances, than browsing ruminants, with the resulting hypothesis that grazers should have larger, heavier chewing muscles than browsers. However, the only investigation of this so far [Axmacher and Hofmann (J Zool 215:463-473, 1988)] did not find differences between ruminant feeding types in the masseter muscle mass of 22 species. Here, we expand the dataset to 48 ruminant species. Regardless of phylogenetic control in the statistical treatment, there was a significant positive correlation of body mass and masseter mass, and also a significant association between percent grass in the natural diet and masseter mass. The results support the concept that ruminant species that ingest more grass have relatively larger masseter muscles, possibly indicating an increased requirement to overcome the resistance of grass forage. The comparative chewing resistance of different forage classes may represent a rewarding field of ecophysiological research.

In the ongoing debate about divergent evolutionary morphophysiological adaptations of grazing and browsing ruminants, the size of the salivary glands has received special attention. Here, we report the most comprehensive dataset on ruminant salivary glands so far, with data on the Glandula parotis (n=62 species), Gl. mandibularis (n=61), Gl. buccalis ventralis (n=44), and Gl. sublingualis (n=30). All four salivary gland complexes showed allometric scaling with body mass (BM); in all cases, the 95% confidence interval for the allometric exponent included 0.75 but did not include 1.0 (linearity); therefore, like other parameters linked to the process of food intake, salivary gland mass appears to be correlated to metabolic body weight (BM0.75), and comparisons of relative salivary gland mass between species should rather be made on the basis of BM0.75 than as a percentage of BM. In the subsequent analyses, the percentage of grass (%grass) in the natural diet was used to characterize the feeding type; the phylogenetic tree used for a controlled statistical evaluation was entirely based on mitochondrial DNA information. Regardless of phylogenetic control in the statistical treatment, there was, for all four gland complexes, a significant positive correlation of BM and gland mass, and a significant negative correlation between %grass in the natural diet and gland mass. If the Gl. parotis was analyzed either for cervid or for bovid species only, the negative correlation of gland mass and %grass was still significant in either case; an inspection of certain ruminant subfamilies, however, suggested that a convergent evolutionary adaptation can only be demonstrated if a sufficient variety of ruminant subfamilies are included in a dataset. The results support the concept that ruminant species that ingest more grass have smaller salivary glands, possibly indicating a reduced requirement for the production of salivary tannin-binding proteins.

The function of the 'third compartment' of the ruminant forestomach, the omasum, has been debated for a long time. To date, it is assumed that its major function is fluid reabsorption. In order to investigate differences in this organ between ruminant feeding types, we first compared macroscopic measurements of the omasa of free-ranging muskoxen Ovibos moschatus [n=6, mean body mass (BM) 207 kg, range 180-221], a grazer, and free-ranging moose Alces alces (n=11, mean BM 291 kg, range 144-418), a strict browser. Despite the similar BM range, omasa of muskoxen contained more ingesta, had a higher empty organ weight, had more third- and fourth-order laminae, and represented a higher proportion of the total forestomach weight. In particular, the surface area of the omasal leaves - the area available for fluid absorption - was significantly larger in muskoxen (10 933 +/- 940 cm(2)) than in moose (2228 +/- 885 cm(2)). In order to test whether the difference in available surface area is a true functional correlate of feeding type, additional data on the omasal laminar surface area were generated for 83 individuals of 19 species. These data were supplemented with data on 13 additional species from the literature. The percentage of grass (%grass) in the natural diet was used to characterize the feeding type; the phylogenetic tree used for a controlled statistical evaluation was entirely based on mitochondrial DNA information. Regardless of phylogenetic control in the statistical treatment, there was a significant positive correlation of both BM and %grass in the natural diet with omasal laminar surface area. The data suggest that certain ruminant species that ingest more grass have larger omasal leaf surface areas, possibly indicating a higher need for water reabsorption distal to the ruminoreticulum, which could be explained as a consequence of the more distinct rumen contents stratification in these species.

Roe deer are seasonal breeders with a short rutting season from mid-July to mid-August. The seasonality of reproductive activity in males is associated with cyclic changes between growth and involution of both testes and the accessory sex glands. This study characterizes morphological and functional parameters of these organs prior to, during and after breeding season in live adult roe deer bucks. Size and morphology of the reproductive tract was monitored monthly by transcutaneous (testes, epididymis) and transrectal (accessory glands) ultrasonography. Semen was collected by electroejaculation. Concentration, motility and morphological integrity of spermatozoa as well as the content of proteins and testosterone in semen plasma were evaluated. Proportions of haploid, diploid and tetraploid cells were estimated by flow cytometry in testicular tissue biopsies. Serum testosterone was measured by enzyme immunoassay. Most parts of the male reproductive tract showed distinct circannual changes in size and texture. These changes were most pronounced in the testes, seminal vesicles, and prostate. All reproductive organs were highly developed during the rut only. The volume of ejaculates, total sperm number and percentages of motile and intact spermatozoa also showed a maximum during this period and corresponded with high proportions of haploid cells in the testis. The highest percentages of tetraploid cells were found in the prerutting period. The production of motile and intact spermatozoa correlated with both the protein content of semen plasma and the concentration of testosterone in semen plasma and blood serum. These results suggest the importance of combined actions of the testes and accessory sex glands and the crucial role of testosterone in facilitating the optimal timing of intensified semen production to ensure sufficient numbers of normal spermatozoa in seasonal breeders.

The variable keratin appendages of the forestomach epithelium on both sides of the reticulo-omasal orifice, emerging in early summer and disappearing towards the winter in the moose Alces alces L. were studied. During the study period, droppings of the free-ranging moose were collected randomly for faecal particle size distribution analysis. The morphological distribution, size and abundance of keratin end caps was recorded in moose that had died of various causes, and it correlated with the particle size distribution observed in faeces. The fusiform keratin filter of the reticulo-omasal orifice seems to reflect the forage quality and use. It is suggested that this filter prevents the passage of non-fragmented foliage and twig fragments of excessive size during the period of rapid and non-selective intake of high-quality mature forage and it disappears before the short growing season of easily available succulent vegetation with a high nutrient content.

The dissection of the heads of three adult roe deer (Capreolus capreolus) provided evidence that this species of the Cervidae, like carnivores, lagomorphs and certain members of the Bovidae, possesses a zygomatic salivary gland. Homology of this salivary gland in ruminants and carnivores is assumed on the basis of its over-all topographic position and the number and course of its excretory ducts. The zygomatic gland of roe deer is a polystomatic gland and amounts to 0.02% of body weight. As in Saiga tatarica and Capricornis crispus (Bovidae), its secretions are predominantly serous. Its slightly alterated position in roe deer (and in bovids) seems to be caused by an evolutionary transformation of orbitotemporal proportions and the formation of a lacrimal bulla, in particular. This is an inflation of the intraorbital portion of the lacrimal bone providing ventral support to the ocular bulb. The lacrimal bulla is a late evolved character and emerges late in ontogeny. Its postnatal development in roe deer is presented. Comparison with the neonatal stage of lacrimal morphology in cattle suggests a similar ontogenetic development in the Bovidae. The adult grysbok (Raphicerus melanotis, Neotraginae, Bovidae) provides a ‘model situation’ of the primitive character state at the beginning of the evolutionary development of a lacrimal bulla. Zygomatic gland, lacrimal bulla and other structural elements of the ruminant orbitotemporal region provide an example of the interlocked evolutionary transformation of a complex system, the morphological result of which is a multi-layered compromise between several structurofunctional constraints.

As short chain fatty acids produced in the forestomach are insufficient to satisfy the energy requirements of the concentrate selecting roe deer (Capreolus capreolus), it is proposed that these animals may have other mechanisms to avoid energy losses due to microbial fermentation. Nutrients bypassing down the ventricular groove (rumen bypass) or ruminal escape of unfermented or partially fermented nutrients may be two alternatives. As metabolic evidence for incomplete fermentation in the forestomach we investigated: (1) the abundance of the sodium-dependent glucose co-transporter (SGLT1) in the duodenum; (2) enzyme activities of maltase, saccharase and α-amylase in duodenal and pancreatic tissue; and (3) the proportion of essential, polyunsaturated fatty acids in depot fat samples from ruminants of different feeding type and — for comparison — from animals with a simple stomach. The high abundance of SGLT1, high enzyme activity and the high proportion of polyunsaturated fatty acids in the concentrate selecting ruminants support the hypothesis of rumen bypass or ruminal escape of nutrients in roe deer and reflect differences in nutrient utilization by ruminants that belong to different feeding types.

The structure and fill of the omasum was investigated in summer and in winter in adult female reindeer living on the polar desert and tundra of the high Arctic archipelago of Svalbard and in sub-Arctic mountain habitats in northern Norway The mean total mass of the omasum in non-lactating adult female Svalbard reindeer was 467 g (0.65 g per 100 g live body mass (BM)) in September and 477 g (1.03 g per 100 g BM) in April. By contrast, the mean mass of the omasum in non-lactating adult reindeer in northern Norway was 534 g (0.83 g per 100 g BM) in September but only 205 g (0.35 g per 100 g BM p<0.05) in late March, owing to a decrease in both tissue mass and the wet mass of the contents of the organ. The mean absorptive surface of the omasum in Svalbard reindeer was 2300 cm2 in September and 2023 cm2 in April. In Norwegian reindeer, by contrast, the absorptive surface area decreased from 2201 cm2 in September to 1181 cm2 (p<0.05) in late March. The marked seasonal decline of omasal tissue and contents in Norwegian reindeer probably results from intake of highly digestible forage plants, including lichens, in winter. Svalbard reindeer, a non-migratory sub-species, survive eating poor quality fibrous vascular plants in winter. The absence of any marked seasonal change in the mass, total absorptive surface area or filling of the omasum in Svalbard reindeer in winter despite a substantial decline in body mass presumably reflects their need to maintain maximum absorption of nutrients, including volatile fatty acids, when feeding on such poorly fermentable forage.

The head of an adult male Takin (Budorcas taxicolor) was dissected with special reference to the larynx. Compared to a typical bovid larynx, the larynx of the male B. taxicolor is clearly enlarged. The enlarged larynx is supported ventrally by the very broad parotidoauriculares muscles as if lying in a hammock. Enlargement of the larynx is mainly due to the great size of the thyroid cartilage, which caudoventrally forms a distinct inflation, the thyroid bulla. and of the cricoid cartilage, the dorsal muscular plate and the arcus of which are strongly extended. In contrast to a typical bovid larynx (as exemplified by cattle) a tough cartilaginous connection between the arytenoid cartilages is found dorsomedially. Both, the muscular and the medial process articulate with the rostral edge of the cricoid cartilage. Compared to the typical situation in bovids the vocal process of the arytenoid cartilage is distinctly extended. With respect to these characters the laryngeal structure of the takin bears a greater resemblance to that of cervids (especially to rot deer) than to that of bovids. The vocal fold consists of two portions which differ in mass and flexibility. Formation of a median synchondrotic connection between the arytenoid cartilages, the double articulation of both arytenoid cartilages with the cricoid cartilage and the long, lever-like vocal process presumably are caused by the great width and mass of the voluminous vocal folds. These make higher demands on the tensor apparatus of the vocal folds than it is the case in a typical bovid larynx with narrow vocal folds. This interpretation is supported by the considerable size of the thyroarytenoid muscle. Male takins vocalize mainly during the mating season, whereas females additionally vocalize to call their young. Calls of the female are as low in frequency as those of the male and cannot be distinguished by ear. From this it may be inferred that the larynx of female takins is enlarged to a similar degree like in males. This was confirmed by the dissection of the larynx of a female neonatus which, basically, has the same structure as that in the male sex. The bellowing calls last for about one second. Frequencies are low and mainly cover the range between 0,1 and 4 kHz. Low frequencies are favourable for a communication on longer distances as they are less attenuated than higher ones. In the sonagram two maxima of frequency of the bellowing call are distinguishable: a) at c. 0,24 kHz; b) at c. 0,41 kHz. The higher one is not a harmonic of the lower maximum suggesting the occurrence of nonlinear oscillation dynamics. The male muskox (Ovibos moschatus) possesses an enlarged larynx very much resembling that of the male takin with respect to its external shape. In convergence to the takin and the muskox, the African water chevrotain (Hyemoschus aquaticus. Tragulidae) probably in both sexes evolved an enlarged larynx which, except for its different proportions, is similar to that of the takin. The frequencies of the vocalizations of the water chevrotain cover the same low range as those of the takin, although the larynx of H. aquaticus reaches only one quarter of the takin's laryngeal size. In the reindeer (Rangifer tarandus). males and females possess a median inflatable sac which obviously functions: as a resonance organ for vocalizations which, likewise, lie in the low frequency range.

Knowledge about the mechanisms of embryonic diapause in European roe deer Capreolus capreolus (Linnaeus, 1758), although first described 140 years ago, is not extensive.

Introduction Based on morphological and physiological differences in their digestive systems H ofmann and S tewart (1972) classified ruminants into three feeding types: concentrate selectors (CS), grass and roughage eaters (GR) and intermediate, opportunistic, mixed feeders (IM). In evolutionary terms CS most closely resemble the original ruminants (H ofmann 1989). They have a relatively small forestomach with large openings between the different parts of the stomach. The retention time of ingesta is short and therefore less suited to optimizing cellulose digestion. They have relatively large saliva glands, producing high volumes of saliva (F ickel et al. 1998), which may facilitate the passage of soluble nutrients down their highly developed ventricular groove. The CS ruminants choose easily digestible forage, especially dicotyledons, which are rich in soluble plant cell contents (PCC, T ixier et al. 1997; D uncan et al. 1998). With evolution, the forestomach of the ruminants became larger relative to body size, and also more subdivided. These evolutionary adaptations optimized the selective retention of feed particles and increased retention time, leading to improvement of cellulose digestion. Large GR are well adapted to digest plant cell walls (PCW), i.e. structural carbohydrates (cellulose); they consequently feed mainly on monocotyledons.Roe deer (Capreolus capreolus) are very small ruminants which select concentrated feed. As energy requirements increase proportional to metabolic body weight (K leiber 1961) roe deer require relatively more energy per unit body mass for maintenance compared to that required by large ruminants (V an S oest 1994; D uncan et al. 1998). Easily digestible forage is not sufficient to satisfy the energy requirements of roe deer. It is proposed therefore that these animals may have other methods of avoiding energy losses due to microbial fermentation, either bypassing nutrients down the ventricular groove (rumen bypass) or by ruminal escape of unfermented and/or partially fermented nutrients (H ofmann 1989). The relatively high content of polyunsaturated fatty acids present in the depot fat of roe deer due to only partial rumen biohydrogenation (M eyer et al. 1998) supports the alternative hypothesis of rumen bypass or ruminal escape. Once reaching the small intestine, partially fermented or bypassed soluble carbohydrates can be further hydrolysed by maltase and sucrase to monosaccharides. These monosaccharides are then available for absorption.The intestinal sodium‐dependent glucose cotransporter (SGLT1) in the brush border membrane (BBM) of the enterocytes is a stable indicator for the presence of sugars in the small intestine. The expression and activity of SGLT1 protein are regulated by the level of sugars reaching the small intestine (F erraris and D iamond 1989). Infusion of d‐glucose into the intestinal lumen of ruminant sheep for 2 h stimulates the expression of SGLT1 which can be detected 4 days later in the newly formed mature enterocytes (S hirazi‐B eechey et al. 1995). When dietary carbohydrate is removed, it takes 3 days before the brush border glucose uptake capacity decreases significantly from control values (F erraris 1994). This reflects the time of physiological cell renewal. In GR, such as sheep, the capacity of the intestine to absorb sugars decreases up to 500‐fold during development (S hirazi‐B eechey et al. 1995). This decrease in the intestinal absorptive capacity is a reflection of the decline in SGLT1 transport activity and protein abundance. It is not due to modifications in the structure of the absorptive surface and cannot be explained in terms of an age‐related process (S hirazi‐B eechey et al. 1991).It has been shown that there is a direct correlation between the level of monosaccharides reaching the lumen of the small intestine and the expression of functional SGLT1 protein (D yer et al. 1997). Therefore in this study we aimed to investigate the presence of functional SGLT1 in the small intestine of roe deer as a stable indicator of the availability of glucose in the luminal content of the small intestine. The presence of SGLT1 protein in the small intestine will support the hypothesis of rumen bypass and/or ruminal escape of nutrients. Furthermore we measured the maltase activities in the BBM, as part of the carbohydrates reach the intestinal lumen as polysaccharides and have to be broken down to monosaccharides before absorption.

Many herbivore species have a diet containing high proportions of polyphenolics, principally lignins and tannins; the latter reduce the conversion of ingested nutrients into biomass and exert systemic toxicity at high levels of intake. It has been proposed that tannin-binding proteins in the saliva might be responsible for minimizing these tannin-related effects by forming soluble protein–tannin complexes. We have developed a method that permits evaluation of the relative tannin-binding properties of salivary proteins at a pH of 8.0–8.5. It is aimed at facilitating both the identification of tannin-binding proteins and the investigation of their relative tannin binding. The principle of the assay is the inhibition of trypsin by tannins and the subsequent reversal of that inhibition when other tannin-binding proteins are added (indirect assay). The method is rapid; large sample numbers may be processed by virtue of its microtiterplate format.

Transrectal ultrasonography was performed on five wild and two captive male African elephants (Loxodonta africana) and four captive male Asian elephants (Elephas maximus) to develop standards for assessment of reproductive health and status. The entire internal urogenital tract was visualized ultrasonographically by using a 3.5 MHz or a 7.5-MHz transducer in combination with a probe extension adapted for elephant anatomy. The findings were verified by postmortem ex situ ultrasound examinations in several individuals of each species. Each part of the internal urogenital tract was sonographically detectable except for the bulbourethral glands and the cranial portion of the ureters and ductus deferentes, which were only observed in situ in the neonate. Each structure visualized was measured and described. The size and morphology of the urogenital structures, especially the accessory glands, were indicative of breeding status, if known. There was a notable difference between African and Asian males in the size and morphology of the prostate gland and a slight difference in the shape of the ampullae. No other structure showed significant species differences. The detection of the location and description of the testes may provide information for modifying present castration procedures. Furthermore, ultrasound examination of the male accessory glands may aid in the identification of potential semen donors for assisted reproduction programs in captive elephants.

Ultrasonography is a noninvasive approach for exploration of morphology, topographic anatomy, and biological processes in non-domestic animals. This imaging technique is still not widely used in the biological sciences. Ultrasonography provides reproducible real time images, cross-sectional imaging of tissues, organ structures, and motion. Images may be used for morphometrics, documentation, and preservation of data. Ultrasound imaging in biological sciences offers new information regarding embryonic development, uterine and ovarian function, muscle development, digestive processes, where conventional experimental and dissection techniques are limited. The use of ultrasonography in different vertebrate taxa is combined with characteristic features for applying this imaging technique due to the individual anatomical structures. Recently, miniaturised transducer technology has allowed the development of microprobes with compact, contoured shapes for access to unexplored anatomical regions in non-domestic animals. Many of the new probes offer colour flow imaging, which is an ideal tool for investigating fluid movements inside a body, e.g., blood flow or ingesta few. Improvements in ultrasound technology and the development of new types of application techniques will certainly advance the progress of this imaging modality in biological sciences. The advantages of this technique favour its more widespread use.

The head of an adult male Takin (Budorcas taxicolor) was dissected with special reference to the salivary glands. In addition, important parts of the facial musculature and the skull, the lacrimal bulla in particular, were investigated. The dissection was carried out on fresh material submerged in water. Consecutive dissection Steps were documented by a series of photographic slides and several proportional drawings. The ratio between the mass of the salivary glands and body mass for the Takin is slightly above the average value for the feeding type of a grass and roughage eater (GR). The morphology of the salivary glands, the small size of the parotid gland in particular, support the classification of the Takin as GR with a tendency to the Intermediate feeding type (GR/IM). (Tendency to LM is further supported by the morphology of the digestive tract.) The ability to cope with a poorly digestible diet during the winter months may have been acquired as a preadaptation before the Takin's restriction to its present refuge habitats. Like the Japanese Serow (Capricornis crispus, Caprinae) and the Sai,oa-Antelope (Saiga tatarica, Saiginae), the Takin possesses an additional compact salivary gland which so far has not been described, neither in the domestic nor in other free ranging ruminants investigated. According to its position as well as to the number and course of its excretory ducts this gland is the homologue of the glandula zygomatica of carnivores. Provided that this is correct the glandula zygomatica appears to be part of the basic plan of ruminants. Apparently it has been retained as a pie siomorphic character in only a few extant species. The partial or complete reduction of this salivary gland in the phylogeny of the ruminants may have been caused by a change of feeding habits. However, the reduction may also result from the increasing separation of the orbita from the temporal fossa narrowing the interspace where the glandula zygomatica is located. The separation serves to protect the ocular bulb from adverse pressures effected by the masticatory musculature occurring particularly in late evolved GR due to constant lateral grinding movements of the lower jaw during extensive rumination for particle size reduction.

In ruminants, different functions have been ascribed to the different salivary glands according to the feeding type. In this context, possible adaptations of salivary functions were investigated regarding the secretion of various proteins by different types of salivary glands. To yield uncontaminated parotid saliva in large quantities, a non-surgical method has been developed. Parotid gland secretions were collected via endoscopic placement of guide wires into each parotid duct, which were subsequently used for placement of collection catheters. Salivary flow was stimulated by intra-glandular administration of the parasympathomimetic compound pilocarpine-hydrochloride into the parotid gland. Mixed saliva (excluding parotid saliva) was collected into sterile tubes by normal outflow during the sampling of parotid saliva. The total flow volume, flow rate and the content of proteins as well as of several ions (Na + , K + , Ca 2+ , inorganic phosphate) of both types of saliva were measured in sheep, fallow deer and roe deer. Roe deer secreted the highest amount of total salivary proteins relative to body mass [mg/kg body mass] and the highest relative volume [ml/10 min/kg body mass], both in parotid and mixed saliva, of all ruminant species examined. Additionally, the protein profile and the tannin-binding properties of parotid and mixed saliva in roe deer were investigated. Parotid saliva bound almost twice as much tannin as mixed saliva, underlining the importance of yielding uncontaminated parotid saliva for tannin-binding studies.