Abstract
Presently, the equid lineage occurs completely outside tropical rainforest environments, which is thought of as the cradle of Perissodactyls and early equid ancestors. The ancestral food of those early equids was based on seeds, fruits, foliage and C3-grasses. The CO2-content of the atmosphere was very high, and C4-grasses had not evolved yet. Zebras and horses are considered to be typical grazers often on C4-grasses, even though Asian equids (and the mountain zebra) have a high proportion of browse in their diet. In comparison to more open environments, present-day tropical forests represent marginal habitats for large ungulates in comparison to more open environments. The suite of traits of large ungulates is not very well adapted to this environment, but equids should not be considered the pinnacle of adaptation to open environments; they retain basal tropical forest traits and lack certain derived open environment traits. Also, equids should not be considered some “non-ruminating ruminant”: their physiology is very different from that of the artiodactyls with which they are often compared; in particular, they are much better at digesting starch and other soluble carbohydrates. We present four storylines on why extant equids may be absent from tropical rainforests: one centred on carbon dioxide, one on chemical plant defence, one on metabolism, and finally a parasitism storyline. Storylines are helpful to envisage how things could have evolved but, of course, do not provide proof.
Access this chapter
Tax calculation will be finalised at checkout
Purchases are for personal use only
References
Anthony, D. W. (1994). The earliest horseback riders and Indo-European origins: New evidence from the steppes. In B. Hänsel & S. Zimmer (Eds.), Die Indogermanen und das Pferd (pp. 185–195). Archaeolingua.
Antonia, E., Merle, R., Stumpff, F., Bollinger, L., Liertz, S., Weber, C., & Heidrun, G. (2021). Evaluation of different blood parameters from endurance horses competing at 160km. Journal of Equine Veterinary Science, 104, 103687.
Bailey, R. C., & Headland, T. N. (1991). The tropical rain forest: is it a productive environment for human foragers? Human Ecology, 19, 261–285.
Bard, J. B. L. (1981). A model for generating aspects of zebra and other mammalian coat patterns. Journal of Theoretical Biology, 93, 363–385.
Barnett, R., Sinding, M. H. S., Vieira, F. G., Mendoza, M. L. Z., Bonnet, M., Araldi, A., Kienast, I., Zambarda, A., Yamaguchi, N., Henschel, P., & Gilbert, M. T. P. (2018). No longer locally extinct? Tracing the origins of a lion (Panthera leo) living in Gabon. Conservation Genetics, 19, 611–618.
Barton, H., & Paz, V. (2007). Subterranean diets in the tropical rainforests of Sarawak, Malaysia. In T. Denham, J. Iriarte, & L. Vrydaghs (Eds.), Rethinking agriculture: archaeological and ethnoarchaeological perspectives (pp. 50–77). Routledge.
Bekhuis, P. D. B. M., de Jong, C., & Prins, H. H. T. (2008). Diet selection and density estimates of forest buffalo in Campo-Ma’an National Park, Cameroon. African Journal of Ecology, 46, 668–675.
Bell, R. H. V. (1971). A grazing ecosystem in Serengeti. Scientific American, 255, 86–93.
Bernardes, C., Sicuro, F. L., Avilla, L. S., & Pinheiro, A. E. P. (2013). Rostral reconstruction of South American hippidiform equids: New anatomical and ecomorphological inferences. Acta Palaeontologica Polonica, 58, 669–678.
Blom, A., Alers, M. P. T., & Barnes, R. F. W. (1990). Gabon. In R. East (Ed.), Antelopes: Global survey & regional action plans (West and Central Africa) (Vol. 3, pp. 113–120). IUCN.
Bodmer, R. E. (1989). Ungulate biomass in relation to feeding strategy within Amazonian forests. Oecologia, 81, 547–550.
Braz, C. U., Rowan, T. N., Schnabel, R. D., & Decker, J. E. (2021). Genome-wide association analyses identify genotype-by-environment interactions of growth traits in Simmental cattle. Scientific Reports, 11, 1–15.
Brøkner, C., Knudsen, K. B., Karaman, I., Eybye, K. L., & Tauson, A. H. (2012). Chemical and physicochemical characterisation of various horse feed ingredients. Animal Feed Science and Technology, 177, 86–97.
Bryant, J. P., Kuropat, P. J., Reichardt, P. B., & Clausen, T. P. (1991). Controls over the allocation of resources by woody plants to chemical antiherbivore defense. In R. T. Palo & C. T. Robbins (Eds.), Plant defenses against mammalian herbivory. CRC Press.
Bryant, J. P., Reichardt, P. B., & Clausen, T. P. (1992). Chemically mediated interactions between woody plants and browsing mammals. Journal of Range Management, 45, 18–24.
Bullimore, S. R., Pagan, J. D., Harris, P. A., Hoekstra, K. E., Roose, K. A., Gardner, S. C., & Geor, R. J. (2000). Carbohydrate supplementation of horses during endurance exercise: Comparison of fructose and glucose. The Journal of Nutrition, 130(7), 1760–1765.
Caro, T. (2005). The adaptive significance of coloration in mammals. BioScience, 55, 125–136.
Caro, T., Argueta, Y., Briolat, E., Bruggink, J., Kasprowsky, M., Lake, J., Mitchell, M., Richardson, S., & How, M. (2019). Benefits of zebra stripes: behaviour of tabanid flies around zebras and horses. PLoS One, 14, e0210831.
Caro, T., Izzo, A., Reiner, R. C., Jr., Walker, H., & Stankowich, T. (2014). The function of zebra stripes. Nature Communications, 5, 3535.
Charles-Dominique, T., Davies, T. J., Hempson, G. P., Bezeng, B. S., Daru, B. H., Kabongo, R. M., Maurin, O., Muasya, A. M., van der Bank, M., & Bond, W. J. (2016). Spiny plants, mammal browsers, and the origin of African savannas. Proceedings of the National Academy of Sciences, 113, E5572–E5579.
Cheeke, P. R., & Dierenfeld, E. S. (2010). Comparative animal nutrition and metabolism. CABI.
Clauss, M., Gehrke, J., Hatt, J. M., Dierenfeld, E. S., Flach, E. J., Hermes, R., Castella, J., Streich, J., & Fickel, J. (2005). Tannin-binding salivary proteins in three captive rhinoceros species. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 140, 67–72.
Clauss, M., & Hummel, J. (2005). The digestive performance of mammalian herbivores: why big may not be that much better. Mammal Review, 35, 174–187.
Clauss, M., Lason, K., Gehrke, J., Lechner-Doll, M., Fickel, J., Grune, T., & Streich, W. J. (2003). Captive roe deer (Capreolus capreolus) select for low amounts of tannic acid but not quebracho: fluctuation of preferences and potential benefits. Comparative Biochemistry and Physiology B, 136, 369–382.
Clauss, M., Steuer, P., Muller, D. W. H., Codron, D., & Hummel, J. (2013). Herbivory and body size: allometries of diet quality and gastrointestinal physiology, and implications for herbivore ecology and dinosaur gigantism. PLoS One, 8, e68714.
Cleaveland, S., Packer, C., Hampson, K., Kaare, M., Kock, R., Craft, M., Lembo, T., Mlengeya, T., & Dobson, A. (2008). The multiple roles of infectious diseases in the Serengeti ecosystem. In A. R. E. Sinclair, C. Packer, S. A. R. Mduma, & J. M. Fryxell (Eds.), Serengeti III: Human impacts on ecosystem dynamics (pp. 209–239). University of Chicago Press.
Coley, P. D. (1983). Herbivory and defensive characteristics of tree species in a lowland tropical forest. Ecological Monographs, 53, 209–234.
Coley, P. D., & Barone, J. A. (1996). Herbivory and plant defenses in tropical forests. Annual Review of Ecology and Systematics, 27, 305–335.
Crile, G., & Quiring, D. P. (1940). A record of the body weight and certain organ and gland weights of 3960 animals. Ohio Journal of Science, 40, 219–259.
Daugirdas, J. T., Levin, N. W., Kotanko, P., Depner, T. A., Kuhlmann, M. K., Chertow, G. M., & Rocco, M. V. (2008). Comparison of proposed alternative methods for rescaling dialysis dose: resting energy expenditure, high metabolic rate organ mass, liver size, and body surface area. Seminars in Dialysis, 21, 377–384.
De Castro, F., & Bolker, B. (2005). Mechanisms of disease-induced extinction. Ecology Letters, 8, 117–126.
Deng, A. Y., & Ménard, A. (2020). Conserved mammalian modularity of quantitative trait loci revealed human functional orthologs in blood pressure control. PLoS One, 15, e0235756.
Ellis, A. D., Longland, A. C., Coenen, M., & Miraglia, N. (2010). Biological basis of behaviour in relation to nutrition and feed intake in horses. EAAP Publication, 128, 53–74.
Estes, R. D. (1991). The behavior guide to African mammals. University of California Press.
Faith, J. T. (2014). Late Pleistocene and Holocene mammal extinctions on continental Africa. Earth-Science Reviews, 128, 105–121.
Fereidouni, S., Freimanis, L., Orynbayev, M., Ribeca, P., Flannery, J., King, D. P., Zuther, S., Beer, M., Höper, D., Kydyrmanov, A., Karamendin, K., & Kock, R. (2019). Mass die-off of saiga antelopes, Kazakhstan, 2015. Emerging Infectious Diseases, 25, 1169–1176.
Gambiza, J. (2001). A primer on savanna ecology. IES Special Report no. 18. UZ, IES.
Geist, V. (1978). Life strategies, human evolution, environmental design: Toward a biological theory of health. Springer.
Geist, V. (1987). On speciation in Ice Age mammals, with special reference to cervids and caprids. Canadian Journal of Zoology, 65, 1067–1084.
Geist, V. (1998). Deer of the World: their evolution, behaviour, and ecology. Stackpole Books.
Gentry, A. W. (1990). Evolution and dispersal of African Bovidae. In G. A. Bubenik & A. B. Bubenik (Eds.), Horns, pronghorns and antlers: Evolution, morphology, physiology, and social significance. Springer.
Geor, R. J. (2010). Digestive strategy and flexibility in horses with reference to dietary carbohydrates. In The impact of nutrition on the health and welfare of horses (pp. 17–28). Wageningen Academic Publishers.
Gessner, J., Buchwald, R., & Wittemyer, G. (2013). Assessing species occurrence and species-specific use patterns of bais (forest clearings) in Central Africa with camera traps. African Journal of Ecology, 52, 59–68.
Gibson, G. (1992). Do tsetse flies “see” zebras? A field study of the visual response of tsetse to striped targets. Physiological Entomology, 17, 141–147.
Gibson, G., & Young, S. (1991). The optics of tsetse fly eyes in relation to their behaviour and ecology. Physiological Entomology, 16, 273–282.
Goachet, A. G., Deschamps, S., Auffret, P., & Julliand, V. (2012). Feeding levels in endurance horses: Practical studies on French riders competing in tests of 130 and 160 km. Journée de la recherche équine, 38, 153–162.
Gordon, I. J., & Prins, H. H. T. (2019). Browsers and grazers drive the dynamics of ecosystems. In I. J. Gordon & H. H. T. Prins (Eds.), The ecology of grazing and browsing II. Ecological studies (Vol. 239, pp. 405–444). Springer.
Gosling, L. M., & Kingdon, J. (2013). Tribe Alcelaphini. In J. Kingdon & M. Hoffman (Eds.), Mammals of Africa, volume VI: Pigs, hippopotamuses, chevrotain, giraffes, deer, and bovids (pp. 488–489). Bloomsbury.
Gould, S. J., & Lewontin, R. C. (1979). The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proceedings of the Royal Society of London. Series B. Biological Sciences, 205, 581–598.
Groot-Bruinderink, G. W. T. A., Lammertsma, D. R., Kramer, K., Wijdeven, S., Baveco, J. M., Kuiters, A. T., Cornelissen, P., Vulink, J. T., Prins, H. H. T., van Wieren, S. E., de Roder, F., & Wigbels, W. (1999). [Dynamic interaction between ungulates and vegetation in the Oostvaardersplassen] (in Dutch: Dynamische interacties tussen hoefdieren en vegetatie in de Oostvaardersplassen). IBN, Report 436 (p. 132).
Guthrie, D. (1990). Frozen fauna of the mammoth steppe; the story of Blue Babe. University of Chicago Press.
Habermann, E., Dias de Oliveira, E. A., Contin, D. R., Delvecchio, G., Viciedo, D. O., de Moraes, M. A., de Mello Prado, R., de Pinho Costa, K. A., Braga, M. R., & Martinez, C. A. (2019). Warming and water deficit impact leaf photosynthesis and decrease forage quality and digestibility of a C4 tropical grass. Physiologia Plantarum, 165, 383–402.
Halsted, M., & Lynch, J. (1996). Phosphorus responses of C3 and C4 species. Journal of Experimental Botany, 47, 497–505.
Harrison, S. (1987). Treefall gaps versus forest understory as environments for a defoliating moth on a tropical forest shrub. Oecologia, 72, 65–68.
Haynes, G. (1995). Pre-Clovis and Clovis Megamammals: a comparison of carcass disturbance, age profiles and other characteristics in light of recent actualistic studies. In E. Johnson (Ed.), Ancient peoples and landscapes (pp. 9–27). Museum of Texas Tech University.
Hempson, G. P., Archibald, S., & Bond, W. J. (2015). A continent-wide assessment of the form and intensity of large mammal herbivory in Africa. Science, 350, 1056–1061.
Hess, T. M., Kronfeld, D. S., Treiber, K. H., Crandell, K. E., Waldron, J. N., Williams, C. A., Staniar, W. B., Lopes, M. A., & Harris, P. A. (2007). Fat adaptation affects insulin sensitivity and elimination of horses during an 80 km endurance ride. Pferdeheilkunde, 23, 241–246.
Hibert, F., Sabatier, D., Andrivot, J., Scotti-Saintagne, C., Gonzalez, S., Prevost, M.-F., Grenand, P., Chave, J., Caron, H., & Richard-Hansen, C. (2011). Botany, genetics and ethnobotany: a crossed investigation on the Elusive Tapir’s Diet in French Guiana. PLoS One, 6(10), e25850.
Hofmann, R. R. (1989). Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system. Oecologia, 78, 443–457.
Hofmann, R. R., & Stewart, D. R. M. (1972). Grazer or browser: a classification based on the stomach-structure and feeding habits of East African ruminants. Mammalia, 36, 226–240.
Hopcraft, J. G. C., Anderson, T. M., Pérez-Vila, S., Mayemba, E., & Olff, H. (2012). Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers. Journal of Animal Ecology, 81, 201–213.
Horváth, G., Pereszlényi, Á., Száz, D., Barta, A., Jánosi, I. M., Gerics, B., & Åkesson, S. (2018). Experimental evidence that stripes do not cool zebras. Scientific Reports, 8, 1–12.
Hughes, A. E., Magor-Elliott, R. S., & Stevens, M. (2015). The role of stripe orientation in target capture success. Frontiers in Zoology, 12(1), 1–12.
Hurst, L. D., Pál, C., & Lercher, M. J. (2004). The evolutionary dynamics of eukaryotic gene order. Nature Reviews Genetics, 5, 299–310.
Ireland, H. M., & Ruxton, G. D. (2017). Zebra stripes: an interspecies signal to facilitate mixed-species herding? Biological Journal of the Linnean Society, 121, 947–952.
Janis, C. (1984). Tapirs as living fossils. In N. Eldredge & S. M. Stanley (Eds.), Living fossils. Casebooks in earth sciences. Springer.
Janis, C. (2007). Artiodactyl paleoecology and evolutionary trends. In D. R. Prothero & S. E. Foss (Eds.), Evolution of artiodactyls. The Johns Hopkins University Press.
Janis, C. (2009). Artiodactyl ‘success’ over perissodactyls in the late Palaeogene unlikely to be related to the carotid rete: a commentary on Mitchell & Lust (2008). Biology Letters, 5(1), 97.
Janis, C. M., & Bernor, R. L. (2019). The evolution of equid monodactyly: a review including a new hypothesis. Frontiers in Ecology and Evolution, 7, 119.
Kawecki, T. J. (2008). Adaptation to marginal habitats. Annual Review of Ecology, Evolution, and Systematics, 39, 321–342.
Kerley, G. I. H., Kowalczyk, R., & Cromsigt, J. P. G. (2012). Conservation implications of the refugee species concept and the European bison: king of the forest or refugee in a marginal habitat? Ecography, 35, 519–529.
Kiltie, R. A. (1984). Seasonality, gestation time, and large mammal extinctions. In P. S. Martin & R. G. Klein (Eds.), Quaternary extinctions (pp. 299–314). University of Arizona Press.
Kingdon, J. (1990). Island Africa. HarperCollins Publishers.
Kingdon, J. (2013). Genus Tragelaphus spiral-horned Antelopes. In J. Kingdon & M. Hoffmann (Eds.), Mammals of Africa, volume VI pigs, hippopotamuses, chevrotain, giraffes, deer and bovids. Bloomsbury.
Klaus-Hügi, C., Klaus, G., & Schmid, B. (1999). Feeding ecology of a large social antelope in the rainforest. Oecologia, 119, 81–90.
Kreulen, D. A. (1985). Lick use by large herbivores: a review of benefits and banes of soil consumption. Mammal Review, 15, 107–123.
Lapierre, H., & Lobley, G. E. (2001). Nitrogen recycling in the ruminant: A review. Journal of Dairy Science, 84, E223–E236.
Larison, B., Harrigan, R. J., Thomassen, H. A., Rubenstein, D. I., Chan-Golston, A. M., Li, E., & Smith, T. B. (2015). How the zebra got its stripes: a problem with too many solutions. Royal Society Open Science. https://doi.org/10.1098/rsos.140452
Law, R. (1976). Horses, firearms, and political power in pre-colonial West Africa. Past & Present, 72, 112–132.
Librado, P., & Orlando, L. (2021). Genomics and the evolutionary history of equids. Annual Review of Animal Biosciences, 9, 81–101.
Loutit, B. D., Louw, G. N., & Seely, M. K. (1987). First approximation of food preferences and the chemical composition of the diet of the desert dwelling black rhinoceros (Diceros bicornis). Madoqua, 15, 35–54.
Lovegrove, B. G. (2012a). The evolution of mammalian body temperature: the Cenozoic supraendothermic pulses. Journal of Comparative Physiology B, 182, 579–589.
Lovegrove, B. G. (2012b). The evolution of endothermy in Cenozoic mammals: a plesiomorphic-apomorphic continuum. Biological Reviews of the Cambridge Philosophical Society, 87, 128–162.
Lovegrove, B. G. (2016). A phenology of the evolution of endothermy in birds and mammals. Biological Reviews of the Cambridge Philosophical Society, 92, 1213–1240.
Lundgren, E. J., Ramp, D., Stromberg, J. C., Wu, J., Nieto, N. C., Sluk, M., Moeller, K. T., & Wallach, A. D. (2021). Equids engineer desert water availability. Science, 6541, 491–495.
MacFadden, B. J. (1993). Fossil horses: Systematics, paleobiology, and evolution of the family Equidae. Cambridge University Press.
Marderstein, A. R., Davenport, E. R., Kulm, S., Van Hout, C. V., Elemento, O., & Clark, A. G. (2021). Leveraging phenotypic variability to identify genetic interactions in human phenotypes. The American Journal of Human Genetics, 108, 49–67.
Markman, A. B., & Wood, K. L. (Eds.). (2009). Tools for innovation: The science behind the practical methods that drive new ideas. Oxford University Press.
Marlowe, F. (2002). Why the Hadza are still hunter-gatherers. In S. Kent (Ed.), Ethnicity, hunter-gatherers, and the “other”: Association or Assimilation in Africa (pp. 247–275). Smithsonian Institution Press.
Martin, R. (2005). Tree-kangaroos of Australia and New Guinea. CSIRO.
McArthur, C., Sanson, G. D., & Beal, A. M. (1995). Salivary proline-rich proteins in mammals: Roles in oral homeostasis and counteracting dietary tannin. Journal of Chemical Ecology, 21, 663–691.
McNaughton, S. J. (1985). Ecology of a grazing ecosystem: the Serengeti. Ecological Monographs, 55, 259–294.
Medici, E. P., Mangini, P. R., Veloso Nunes, A. L., & Vaz Ferreira, J. R. (2008). In M. E. Fowler & Z. S. Cubas (Eds.), Order Perissodactyla, family Tapiridae (Tapirs). Biology, medicine, and surgery of South American wild animals (pp. 363–376). Iowa State University Press.
Merediz, E. F. C., Dyer, J., Salmon, K. S. H., & Shirazi-Beechey, S. P. (2004). Molecular characterisation of fructose transport in equine small intestine. Equine Veterinary Journal, 36(6), 532–538.
Meyer, K., Hummel, J., & Clauss, M. (2010). The relationship between forage cell wall content and voluntary food intake in mammalian herbivores. Mammal Review, 40(3), 221–245.
Mishra, C., Khanyari, M., Prins, H. H. T., & Suryawanshi, K. R. (2019). Community dynamics of browsing and grazing ungulates. In I. J. Gordon & H. H. T. Prins (Eds.), The ecology of grazing and browsing II (pp. 181–196). Springer.
Mitchell, D., Maloney, S. K., Jessen, C., Laburn, H. P., Kamerman, P. R., Mitchell, G., & Fuller, A. (2002). Adaptive heterothermy and selective brain cooling in arid-zone mammals. Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 131, 571–585.
Mitchell, G., & Lust, A. (2008). The carotid rete and artiodactyls success. Biology Letters, 4, 415–418.
Mkhize, N. R., Heitkӧnig, I. M. A., Scogings, P. F., Dziba, L. E., Prins, H. H. T., & de Boer, W. F. (2018). Effects of condensed tannins on live weight, faecal nitrogen and blood metabolites of free-ranging female goats in a semi-arid African savanna. Small Ruminant Research, 166, 28–34.
Mkhize, N. R., Heitkönig, I. M. A., Scogings, P. F., Hattas, D., Dziba, I. E., Prins, H. H. T., & De Boer, W. F. (2016). Supplemental nutrients increase the consumption of chemically defended shrubs by free-ranging herbivores. Agriculture, Ecosystems & Environments, 235, 119–126.
Montenegro, O. L. (1998). The behavior of Lowland Tapir (Tapirus terrestris) at a natural mineral lick in the Peruvian Amazon. MSc thesis, University of Florida.
Naude, T. W., Wood, P. A., & Foggin, D. C. (1997). Suspected calcium oxalate raphide irritation in a black rhinoceros (Diceros bicornis) due to ingestion of Xanthosoma mafaffa. Journal of the South African Veterinary Association, 68, 2–2.
Nieschling, H. (1935). Digestibility of raw carrots by draught horses. Zeitschrift für Züchtung, Reihe B, 32, 331–348.
NRC National Research Council. (1989). Nutrient requirements of horses (5th revised ed). National Academy Press.
Osborn, H. F. (1910). The age of mammals in Europe, Asia, and North America. Macmillan.
Ostfeld, R. S., Keesing, F., & Eviner, V. T. (2008). Infectious disease ecology: the effects of ecosystems on disease and of disease on ecosystems. Princeton University Press.
Pearson, P. N., & Palmer, M. R. (2000). Atmospheric carbon dioxide concentrations over the past 60 million years. Nature, 406, 695–699.
Pretorius, Y., de Boer, W. F., Kortekaas, K., van Wijngaarden, M., Grant, R. C., Kohi, E. M., & Prins, H. H. T. (2016). Why elephant have trunks and giraffe long tongues: how plants shape large herbivore mouth morphology. Acta Zoologica, 97, 246–254.
Preuss, S., & Pikaard, C. S. (2007). rRNA gene silencing and nucleolar dominance: insights into a chromosome-scale epigenetic on/off switch. Biochimica et Biophysica Acta (BBA)-Gene Structure and Expression, 1769, 383–392.
Prins, H. H. T. (1996). Behaviour and ecology of the African Buffalo: Social inequality and decision making. Chapman & Hall.
Prins, H. H. T. (2016). Interspecific resource competition in antelopes: search for evidence. In J. Bro-Jorgensen & D. P. Mallon (Eds.), Antelope conservation: from diagnosis to action, Conservation science and practice series (pp. 51–77). Wiley Blackwell.
Prins, H. H. T., & Olff, H. (1998). Species richness of African grazer assemblages: Towards a functional explanation. In D. Newbery, H. H. T. Prins, & N. D. Brown (Eds.), Dynamics of tropical communities (pp. 449–490). Blackwell.
Prins, H. H. T., Ottenburghs, J., & van Hooft, P. (2023). The environments of the African buffalo, with different selection forces in different habitats. In A. Caron, D. Cornélis, P. Chardonnet, & H. H. T. Prins (Eds.), Ecology and management of the African Buffalo. Cambridge University Press.
Prins, H. H. T., & Reitsma, J. M. (1989). Mammalian biomass in an African equatorial rain forest. Journal of Animal Ecology, 58, 851–861.
Prins, H. H. T., & Sinclair, A. R. E. (2013). Syncerus caffer African buffalo. In J. Kingdon & M. Hoffmann (Eds.), Mammals of Africa, volume VI pigs, hippopotamuses, chevrotain, giraffes, deer and bovids. Bloomsbury.
Probst, J. R., & Hayes, J. P. (1987). Pairing success of Kirtland’s Warblers in marginal vs. suitable habitat. The Auk, 104, 234–241.
Prokkola, J. M., Nikinmaa, M., Lewis, M., Anttila, K., Kanerva, M., Ikkala, K., & Leder, E. H. (2018). Cold temperature represses daily rhythms in the liver transcriptome of a stenothermal teleost under decreasing day length. Journal of Experimental Biology, 221(5), jeb170670.
Prothero, D. R., & Schoch, R. M. (1989). The evolution of perissodactyls. Oxford University Press.
Prothero, D. R., & Schoch, R. M. (2002). Horns, Tusk and Flippers: the evolution of hoofed mammals. The Johns Hopkins University Press.
Putz, F. E., & Holbrook, N. M. (1988). Tropical rain-forest images. In J. S. Denslow & C. Padoch (Eds.), People of the tropical rain forest (pp. 37–52). University of California Press.
Reynolds, C. K., & Kristensen, N. B. (2008). Nitrogen recycling through the gut and the nitrogen economy of ruminants: an asynchronous symbiosis. Journal of Animal Science, 86, E293–E305.
Richards, L., & Coley, P. D. (2007). Seasonal and habitat differences affect the impact of food and predation on herbivores: a comparison between gaps and understory of a tropical forest. Oikos, 116, 31–40.
Ritchie, M. E., & Olff, H. (1999). Spatial scaling laws yield a synthetic theory of biodiversity. Nature, 400, 557–560.
Robbins, C. T. (1993). Wildlife feeding and nutrition (2nd ed.). Academic Press.
Salas, L. A., & Fuller, T. K. (1996). Diet of the lowland tapir (Tapirus terrestris L.) in the Tabaro River valley, southern Venezuela. Canadian Journal of Zoology, 74(8), 1444–1451.
Saltzman, E. S., Petit, J. R., Jouzel, J., Raynaud, D., Barkov, N. I., Barnola, J. M., Basile, I., Bender, M., Chappellaz, J., Davis, J., Delaygue, G., Delmotte, M., Kotlyakov, V. M., Legrand, M., Lipenkov, V., Lorius, C., Pépin, L., Ritz, C., & Stievenard, M. (1999). 420,000 years of climate and atmospheric history revealed by the Vostok deep Antarctic ice core. Nature, 399, 429–436.
Sandoval-Castellanos, E., Wutke, S., Gonzalez-Salazar, C., & Ludwig, A. (2017). Coat colour adaptation of post-glacial horses to increasing forest vegetation. Nature Ecology & Evolution, 1, 1816–1819.
Schefuß, E., Schouten, S., Jansen, J. F., & Damsté, J. S. S. (2003). African vegetation controlled by tropical sea surface temperatures in the mid-Pleistocene period. Nature, 422, 418–421.
Schieltz, J. M., & Rubenstein, D. I. (2016). Evidence based review: positive versus negative effects of livestock grazing on wildlife. What do we really know? Environmental Research Letters, 11, 113003.
Schmitz, O. J., & Nudds, T. D. (1994). Parasite-mediated competition in deer and moose: How strong is the effect of meningeal worm on moose? Ecological Applications, 4, 91–103.
Schroder, B. (2021). Methods to improve forage quality for mammalian herbivores in nutrient poor savannas. PhD thesis, Wageningen University.
Searle, K. R., & Shipley, L. A. (2008). The comparative feeding behaviour of large browsing and grazing herbivores. In I. J. Gordon & H. H. T. Prins (Eds.), The ecology of browsing and grazing. Ecological studies (Vol. 195, pp. 117–148). Springer.
Sinatra, G. M., & Lombardi, D. (2020). Evaluating sources of scientific evidence and claims in the post-truth era may require reappraising plausibility judgments. Educational Psychologist, 55, 120–131.
Sinclair, A. R. E. (1979). The eruption of the ruminants. In A. R. E. Sinclair & M. Norton-Griffiths (Eds.), Serengeti: dynamics of an ecosystem (pp. 82–103). University of Chicago Press.
Solounias, N., & Moelleken, S. M. (1992). Tooth microwear analysis of Eotragus sansaniensis (Mammalia: Ruminantia), one of the oldest known bovids. Journal of Vertebrate Paleontology, 12(1), 113–121.
Solounias, N., & Semprebon, G. (2002). Advances in the reconstruction of ungulate ecomorphology with application to early fossil equids. American Museum Novitates, 2002(3366), 1–49.
Soulé, M. (1973). The epistasis cycle: a theory of marginal populations. Annual Review of Ecology and Systematics, 4, 165–187.
Stahl, W. R. (1962). Similarity and dimensional methods in biology. Science, 13, 205–212.
Stitt, M., & Krapp, A. (1999). The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background. Plant, Cell & Environment, 22, 583–621.
Sutherland, B. J., Prokkola, J. M., Audet, C., & Bernatchez, L. (2019). Sex-specific co-expression networks and sex-biased gene expression in the salmonid Brook Charr Salvelinus fontinalis. G3: Genes, Genomes, Genetics, 9, 955–968.
Sutherland, H. G., Kearns, M., Morgan, H. D., Headley, A. P., Morris, C., Martin, D. I., & Whitelaw, E. (2000). Reactivation of heritably silenced gene expression in mice. Mammalian Genome, 11, 347–355.
Talon, I., Janiszewski, A., Chappell, J., Vanheer, L., & Pasque, V. (2019). Recent advances in understanding the reversal of gene silencing during X chromosome reactivation. Frontiers in Cell and Developmental Biology, 7, 169.
Tan, Z., & Murphy, M. R. (2004). Ammonia production, ammonia absorption, and urea recycling in ruminants: a review. Journal of Animal and Feed Sciences, 13, 389–404.
Taylor, C. R. (1966). The vascularity and possible thermoregulatory function of the horns in goats. Physiological Zoology, 39, 127–139.
Thewissen, J. G. M. (2014). The Walking Whales: from land to water in eight million years. University of California Press.
Tomlinson, K. W., van Langevelde, F., Ward, D., Prins, H. H. T., de Bie, S., Vosman, B., Sampaio, E. V. S. B., & Sterck, F. J. (2016). Defence against vertebrate herbivores trades off into architectural and low nutrient strategies amongst savanna Fabaceae species. Oikos, 125, 126–136.
Tucker, M. A., Böhning-Gaese, K., Fagan, W. F., Fryxell, J. M., Van, M. B., Alberts, S. C., Ali, A. H., Allen, A. M., Attias, N., Avgar, T., Bartlam-Brooks, H., Bayarbaatar, B., Belant, J. L., Bertassoni, A., Beyer, D., Bidner, L., van, B. F. M., Blake, S., Blaum, N., & Mueller, T. (2018). Moving in the anthropocene: global reductions in terrestrial mammalian movements. Science, 359, 466–469.
Turner, N. J., Łuczaj, Ł. J., Migliorini, P., Pieroni, A., Dreon, A. L., Sacchetti, L. E., & Paoletti, M. G. (2011). Edible and tended wild plants, traditional ecological knowledge and agroecology. Critical Reviews in Plant Sciences, 30, 198–225.
Valenzuela-Sánchez, A., Wilber, M. Q., Canessa, S., Bacigalupe, L. D., Muths, E., Schmidt, B. R., Cunningham, A. A., Ozgul, A., Johnson, P. T. J., & Cayuela, H. (2021). Why disease ecology needs life-history theory: a host perspective. Ecology Letters, 24, 876–890.
Van der Zon, A. P. (1992). Graminées du Cameroun. Ph.D. thesis, University of Wageningen.
Vanleeuwe, H., & Gautier-Hion, A. (1998). Forest elephant paths and movements at the Odzala National Park, Congo: the role of clearings and Marantaceae forests. African Journal of Ecology, 36, 174–182.
Van Soest, P. J. (1982). Nutritional Ecology of the Ruminant: ruminant metabolism, nutritional strategies, the cellulolytic fermentation and the chemistry of forages and plant fibers. O & B Books.
Veldhuis, M. P., Hofmeester, T. R., Balme, G., Druce, D. J., Pitman, R. T., & Cromsigt, J. P. G. M. (2020). Predation risk constrains herbivores’ adaptive capacity to warming. Nature Ecology & Evolution, 4(8), 1069–1074.
Venter, J. A., Vermeulen, M. M., & Brooke, C. F. (2019). Feeding ecology of large browsing and grazing herbivores. In I. J. Gordon & H. H. T. Prins (Eds.), The ecology of browsing and grazing II. Ecological studies (Vol. 239, pp. 127–153). Springer.
Vereshchagin, N. K. (1974). The mammoth “cemeteries” of north-east Siberia. Polar Record, 17, 3–12.
Vilà-Cabrera, A., Premoli, A. C., & Jump, A. S. (2019). Refining predictions of population decline at species’ rear edges. Global Change Biology, 25, 1549–1560.
Virot, E., Ma, G., Clanet, C., & Jung, S. (2017). Physics of chewing in terrestrial mammals. Scientific Reports, 7, 43967.
Wallace, D. R. (2004). Beasts of Eden: walking whales, dawn horses, and other enigmas of mammal evolution. University of California Press.
Walton, D. N. (1988). Burden of proof. Argumentation, 2, 233–254.
Walton, D. N. (2001). Abductive, presumptive and plausible arguments. Informal Logic, 21, 141–169.
Wand, S. J., Midgley, G. F., Jones, M. H., & Curtis, P. S. (1999). Responses of wild C4 and C3 grass (Poaceae) species to elevated atmospheric CO2 concentration: a meta-analytic test of current theories and perceptions. Global Change Biology, 5, 723–741.
Ward, D., Schmitt, M. H., & Shrader, A. M. (2020). Are there phylogenetic differences in salivary tannin-binding proteins between browsers and grazers, and ruminants and hindgut fermenters? Ecology and Evolution, 10, 10426–10439.
Weckel, M., Giuliano, W., & Silver, S. (2006). Jaguar (Panthera onca) feeding ecology: distribution of predator and prey through time and space. Journal of Zoology, 270(1), 25–30.
Weigelt, J. (1989). Modes of death. In J. Weigelt (Ed.), Recent vertebrate carcasses and their paleobiological implications (pp. 27–78). University of Chicago Press.
Wenxuan, X. U., Weikang, Y. A. N. G., & Jianfang, Q. I. A. O. (2009). Food habits of Kulan (Equus hemionus hemionus) in Kalamaili mountain nature reserve, Xinjiang, China. Acta Theriologica Sinica, 29, 427–431.
Withers, P. C., Cooper, C. E., Maloney, S. K., Bozinovic, F., & Cruz-Neto, A. P. (2016). Ecological and environmental physiology of mammals. Oxford University Press.
Zhang, H., Yan, X., Zhao, Z., & Ji, L. (2013). Analysis of the polysaccharides from Urtica angustifolia and their anti-fatigue activity. African Journal of Pharmacy and Pharmacology, 7, 1438–1447.
Author information
Authors and Affiliations
Corresponding author
Editor information
Editors and Affiliations
Rights and permissions
Copyright information
© 2023 The Author(s), under exclusive license to Springer Nature Switzerland AG
About this chapter
Cite this chapter
de Jong, J.F., Prins, H.H.T. (2023). Why There Are No Modern Equids Living in Tropical Lowland Rainforests. In: Prins, H.H.T., Gordon, I.J. (eds) The Equids. Fascinating Life Sciences. Springer, Cham. https://doi.org/10.1007/978-3-031-27144-1_4
Download citation
DOI: https://doi.org/10.1007/978-3-031-27144-1_4
Published:
Publisher Name: Springer, Cham
Print ISBN: 978-3-031-27143-4
Online ISBN: 978-3-031-27144-1
eBook Packages: Biomedical and Life SciencesBiomedical and Life Sciences (R0)