NDF digestibility: Sheep, Horse and Black Rhinoceros 4 sub-species • • • • South-Central (Diceros bicornis minor) South-Western (Diceros bicornis bicornis) East-African (Diceros bicornis michael) West-African (Diceros bicornis longpipes) • Diceros Greek (Di= two Ceros= horn) and Latin Bicornis (Bi= two Cornis= horn) • So the southwestern black rhino is a two horn two horn two horn…hmm Physical Characteristics • Black rhinos attain a weight of 2,100–2,900 lb (950– 1,300 kg); shoulder height of 56–63 in (143–160 cm); and head and body length of 112–120 in (286–305 cm). Male and female are similar in size. • Despite name, the skin is gray to brownish gray in color, and devoid of hairs. • Horns are made of Keratin. Anterior horn is 16.5–54 in (42–138 cm) in length, the posterior one 8–20 in (20–50 cm). • Have a saddle-backed appearance, rounded ears, and tend to hold the head high, except when feeding on low vegetation. Behavior • Largely solitary, although groups of three to five animals may occasionally form. • A cow and her calf comprise the basic social unit, and adult males are solitary, except when courting a female • When adult males meet, a complex bull ceremony may take place, involving stiff-legged scraping, imposing postures, and short charges sometimes accompanied by screaming groans. • Females are not territorial, but males will tolerate submissive intruder males. Behavior • Males mark the environment with long drag marks made by the legs, spray urine over bushes or other objects, and deposit feces on dung-heaps. • Females use the same dung-heaps and animals of both sexes scatter their droppings with backwardly directed kicks. Distribution The Black Rhinoceros once roamed the lower half of Africa in hundreds of thousands. Today it survives in pockets primarily in Zimbabwe, South Africa, Kenya, Namibia and Tanzania. Feeding Ecology and Diet • Live primarily on grasslands, savannahs, and tropical bushland habitats. • Predominantly low-level browsers, feeding on small saplings and shrubs under 5 ft (1.5 m) in height as well as a variety of herbs, thorny wood, fruit and occasionally small amounts of grass. They often browse in the morning and evening. • Can eat up to 220 different species of plants. • Can live up to 5 days without water during droughts. • Acacia spp. are especially favored, as are various species of Euphorbiaceae, including succulent forms with milky sap reputed to be poisonous. • The prehensile upper lip is used to pull twigs into the mouth, which are then bitten off with the cheek teeth. They crop branch tips up to 0.4 in (10 mm) in thickness and 4–9.8 in (100–250 mm) in length. The horns may be used to bend or break stems to reach higher branches. Bark may also be stripped from certain trees. Diet Composition of Non-Captive Black Rhinoceroses Shrubs/Trees Herbs Grass Source 87-95 5-13 0 Joubert 1971 54-81 18-41 0 Mukinya 1977 81-94 6-19 0 Hall-martin 1982 47-93 5-51 0 Oloo 1994 93-95 3-5 1 Atkinson 1995 56-76 1-11 0-1 Pole 1995 69 31 0 Henning 2001 Digestion Although all rhinos are hind-gut fermenters like rabbits and horses… The fact that Black Rhinos are browsers is very important! – Fundamental difference between fermentation characteristics of browse and grass • Browse has a faster fermentation rate and soon reaches its maximum energy release • Grass has a slow fermentation rate and still yields energy after a longer period of time Clauss et al 2006 Digestion • White Rhino – Squared off upper lip used to “crop” grass – Grazes on savannah • Black Rhino – Prehensile upper lip for browsing – Consumes bushes and shrubs in forest Comparative Anatomy: Black Rhino •The stomach is simple, and the cecum and colon in the hind-gut serve as the main sites of fermentation •Short, wide colon •Gradual transition from colon ascendens to transversum Clauss et al 2003, 2005a Comparative Anatomy: Horse • Monogastric herbivore with extensive post-gastric fermentation •Simple stomach incapable of utilization of forage-based (high fiber) diets •Abrupt shift from colon ascendens to colon transversum Comparative Anatomy: Sheep •Ruminant herbivore with extensive pre-gastric fermentation •Highly developed sacculated stomach capable of extensive and effective utilization of forage-based (high fiber) diets •Extensive fermentation before primary sites of digestion and absorption Comparative Anatomy: length of GIT (in meters) Horse Black Rhino S. Intestine 26.7 (76%) 12.0 (66%) Caecum Colon total DFC total Total GIT 0.7 (5%) 4.9 (28%) 3.7 (20%) 18.5 1.0 (3%) 7.0 (20%) 4.5 (13%) 35.0 Frewein et al (1999) Stevens/Hume (1995) Comparative Anatomy • There are two ways to increase passage time in a tubular system: to shorten the length or to increase diameter • Body length does not always mean longer GIT (horse and tapir longer than an elephant!) • Browsing Rhino’s have a short and wide ceacum (46-91 cm) compared to horses (15-18 cm) Clauss et al 2003 Comparative Anatomy: Average Mean Retention Times 50 45 40 35 MRT (h) 30 25 20 15 10 5 0 Sheep A longer retention time= Slower passage rate Horse B. Rhino Clauss et al 2005a, Illius and Gordon 1995 Clinical Study: Digestion Coefficients for the Black Rhinoceros (Clauss et al 2006) • Objective of this study was to determine if horses were appropriate models for nutrient recommendations for the Black Rhinoceros • Current practice applies the equine model to grazing hind gut fermenters such as the white rhinoceros • It was unknown prior to this study if browsing behavior would reduce digestion coefficients, making the horse a poor model • Did expect that browsers would display a shorter ingesta retention time and lower digestion coefficients as compared to grazers when on comparable diets Clinical Study: Digestion Coefficients for the Black Rhinoceros Materials and Methods: •Eight rhinos were used, selected from 3 zoos. Animals were kept separate during the trial to allow for collection of feces and recording of dietary intake. •Protocols developed for domestic sheep (Mason and Frederickson 1979) were used to evaluate nutritional components including dry matter (DM) and Neutral Detergent Fiber (NDF). •Study compared the digestibility of rhinos to horses using previously published studies. •Additional data on browsing and grazing rhinos was also included from published and unpublished studies. Clinical Study Materials and Methods: • The diets varied in total make-up but included some % of the following: lucerne hay, grass hay, browse, concentrates, fruits and vegetables • Main concern was a crude fiber content that averaged 25 and crude protein averaging 15. Clinical Study: Digestion Coefficients for the Black Rhinoceros Results: When compared with horses from previously conducted studies (on similar rations), the black rhinoceros achieved lower digestion coefficients Clauss Study Results: Horses outperform Rhinos Clinical Study: Digestion Coefficients for the Black Rhinoceros Potential reasons: • Comparatively shorter ingesta retention times in the black rhino (Clauss et al 2005b) •Among wild ruminant species, it has been demonstrated that browsing species achieve lower coefficients than grazing species (Iason and Van Weiren 1999) •Hackenberger (1987) demonstrated that the African elephant which is thought to be adapted to a diet with more browse material than its Asian counterpart also displays shorter retention times and decreased digestion of grass hay Don’t take our word for it… Browsers vs. Grazers Why? • The digestive strategy of long retention times to increase digestion coefficients is an evolutionary adaptation one would expect in grazing species- whether hind gut or foregut fermenters – Grass has a slow fermentative rate and still gives energy even after a long period of time The why, continued • Shorter retention times are expected in browsing species – browse has a faster fermentation rate and soon reaches its maximum energy release – So if you gave a browsing rhino Alfalfa, you’d expect that it wouldn’t reach its full energy potential because its not fermented long enough. You’d also expect that he’d want to eat a lot more. So, the Black Rhino comes to NUT 115 and eats alfalfa pellets… Real and Expected Digestibility order Sheep 63.7 DM/43.9 NDF Horse 58.6 DM/27.2 NDF Rhino ~53 DM/~26 NDF FYI: More on the Vit E Question • • • • • • Problem: Rhinoceros accumulate body stores of iron over their lifetime that may be associated with numerous maladies: ie captive black rhinoceroses have a history of disease and death characterized by hemolytic anemia and mucocutaneous ulceration Current trends: Iron and copper supplements have been routinely given to captive rhinos. In the absence of species specific data, nutrient levels are based on work performed with domestic livestock. Potential Causes: Many nutrients listed as possible causes in the syndromes may be linked in some way to each other Specific nutrients of concern: ferric iron antagonist to vitamin E, interactions between absorption of dietary iron and vitamin E, and linkages between copper and iron absorption/functioning. Wild Rhino Diet: Natural browse of black rhinoceros contains secondary plant compounds including tannins that may decrease mineral absorption. It is possible that these chemicals may bind excess dietary iron in the natural diet. Commercial hays contain tannins, but at lower levels… and are fed to captive rhinos (budget, absence of data, etc) Diets without tannins are hypothesized to cause increased iron absorption and an insufficient antioxidant status, and therefore to contribute to disease symptoms observed in captive black rhinos Current study at the Brookfield Zoo (Illinois): "Dietary Iron Absorption and the Role of Tannins in Eastern (Diceros bicornis michaeli) and Southern Black Rhino (Diceros bicornis minor), a Comparison" Website: http://www.sosrhino.org/research/black_rhino_nutrition2.php last acessed 3/13/06 References CLAUSS M; CASTELL J.C; KIENZLE E; DIERENFELD E.S; FLACH E.J; BEHLERT O; ORTMAN S; STREICH W.J; HUMMEL J; HATT J.M (2006) Digestion coefficients achieved by the black rhinoceros (diceros bicornis), a large browsing hind gut fermenter. Journal of Animal Physiology and Animal Nutrition 90: 325-334. OFTEDAHL, O., BAER, D.J., AND ALLEN, M.E. (1996). The feeding and nutrition of herbivores. In: Kleiman, D.G., Allen, M., Thompson, K.V., and Lumpkin, S. (eds), Wild Mammals in Captivity: Principles and Techniques. University of Chicago Press, Chicago, IL. 129-138. GHEBREMESKEL, K.;WILLIAMS, G.; BRETT, R.A.; BUREK, R.; HARBIGE, L.S (1991) Nutrient composition of plants most favoured by black rhinoceros (Diceros bicornis) in the wild. Comparative Physiology 98: 529– 534. OWEN-SMITH, N. (1975) The social ethology of the white rhinoceros (Ceratotherium simum). Z. Tierpsychology 38: 337. HOFMANN, R.R. (1989) Evolutionary steps of ecophysiological adaptation and diversification of ruminants: A comparative view of their digestive system. Oecologia 78:443-457. CLAUSS M; FROESCHLE T; CASTELL J; HUMMEL J; HATT J-M; ORTMANN S; STREICH W.J (2005) Fluid particle retention times in the black rhinoceros. Acta Theriologica 50: 367-376. CLAUSS M; FREY R; KIEFER B; LECHNER-DOLL M; LOEHLEIN W; POLSTER C; ROSSNER G.E; STREICH W.J (2003) The maximum attainable body size of herbivorous mammals: constraints on forgut, and adaptations of hindgut fermenters. Oecologia 136: 14-27 VAN SOEST, P (1994) Nutritional ecology of the ruminant, 2nd ed. Cornell University Press, Ithaca, NY CLAUSS M; POLSTER C; KIENZLE E; WIESNER H; BAUMGARTNER K; VON HOUWALD F; ORTMANN S; STREICH W.J; DIERENFELD E.S (2005) Studies on digestive physiology and feed digestibilities in captive Indian rhinoceros (Rhinoceros unicornis) Journal of Animal Physiology and Animal Nutrition 89: 229–237. HACKENBERGER M.K., 1987: Diet digestibilities and ingesta transmit times of captive Asian and African elephants. MS thesis, University of Guelph, Canada, pp. 1-115. IASON G.R; VAN WIEREN S.E, 1999: Adaptations of mammalian herbivores to low quality forage. In: H. Olff, V.K Brown, R.H Drent (eds.), Herbivores, Plants and Predators. 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