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Large Herbivores Ecosystem Functions and the Effects of Extinction

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Large Herbivores Ecosystem Functions and the Effects of Extinction Large Herbivores Ecosystem Functions and the Effects of Extinction Angela Harvey IBD 4 Final Review Paper Abstract Large herbivores represent some of the most iconic animals on the planet. These creatures are frequently keystone species as well as flagship species for their environments. Unfortunately, many of them are in danger of extinction in those same habitats. The biomes and ecoregions large grazers and browsers inhabit cover six of the continents, and include grasslands, forests, and shrublan ds. Since many large grazers are keystone species, the services they provide are critical to the health of the ecosystem. The loss of these mammals through extinction would have cascading effects on the areas they inhabit. One order of large herbivores, th e Perissodactyla, are uniquely situated to illustrate herbivory and its function. Since most of the large herbivores are grazers, at least part of the time, grassland ecosystems are used as case studies to highlight the services provided. For background on extinction and its consequences, historical studies of the Pleistocene - Holocene transition are investigated. Introduction Large herbivores represent some of the most iconic animals on the planet, such as the African elephant ( Loxodonta africana ) and the white rhinoceros ( Ceratotherium simum) . These creatures are frequently keystone species as well as flagship species for their environments (Howland et al., 2014; Blair, Nippert, & Briggs, 2014). Unfortunately, many of them are in danger of extinction in t hose same habitats. Approximately 60% of the 74 largest land herbivores are listed as threatened, including all 10 of the largest of Hippopotamidae, Elephantidae, Hominidae, and Tapiridae families as well as 15 of the 20 species of the Suidae, Equidae, Rhi nocerotidae, and Camelidae families (Ripple et al., 2015). Today, there are eight megafauna species (weighing over 1000 kg), a significantly lower number than were present in the late Pleistocene (~42). Seven of these eight are threatened and four are crit ically endangered, with the only species showing improvement being the southern white rhinoceros ( C. simum ) (Ripple et al., 2015), which has increased from fewer than 100 individuals at the turn of the last century to approximately 20,000 today after an i ntensive effort to save the species. The recent increase in poaching may yet place the white rhinoceros ( C. simum ) in company with its other megafauna cousins, though. (Cromsigt & Te Beest, 2014; Sodhi, Brook, & Bradshaw, 2009; Weigl & Knowles, 2014). In a ddition, the ranges of the 74 species of large herbivores is shrinking. Of the 25 species that have had range contraction estimates, they are currently, on average, occupying only 19% of their historical ranges (See Fig. 1) (Ripple et al., 2015). Fig. 1 - “Range contractions over time for three iconic African herbivores. African elephant. ( L. africana ), ca. 1600 versus 2008, common hippopotamus ( Hippopotamus amphibius ), ca. 1959 versus 2008, and black rhinoceros ( Diceros bicornis ), ca. 1700 versus 1987. The historical ranges are in blue, whereas the most recent ranges are represented by darker - colored polygons. For security purposes, the most recent black rhinoceros range polygons (1987) have been moved by random directions” (Ripple et al., 2015, p. 4) T he biomes and ecoregions large grazers and browsers inhabit cover six of the seven continents, including tropical and subtropical moist broadleaf forest, grasslands, shrublands, savannas, mangroves, and other forest or grassy types (See Fig. 2, Ripple et a l., 2015). Since many large grazers are keystone species, the services they provide are critical to the health of the ecosystem. The loss of these mammals through extinction would have cascading effects on the areas they inhabit. Large herbivores shape the landscape through grazing or browsing, trampling, wallowing, and other actions. They can affect other species directly and indirectly through the food web, and they modify abiotic processes by altering nutrient cycles, the properties of the soil, and fire regimes. There is an economic connection to humans, too, when the iconic species of a given territory attracts tourism, creating an economic web around the large herbivores and their environments. These processes cannot be assumed by small herbivores, thu s their ecosystem services are essential (Ripple et al., 2015; Blair et al., 2014; Austrheim, Speed, Martinsen, Mulder, & Mysterud, 2014; Howland et al., 2014; Gill, 2015). Fig. 2 - “Large herbivore total species richness (A) and threatened (B) at the ec oregion level. Ecoregion lists for each species were obtained using the IUCN Red List species range maps and are based on the ecoregions whe re each species is native and currently present”. (Ripple et al., 2015, p. 2) In light of the fragile state of so m any large herbivores, a review of current literature is necessary in understanding the role grazing and browsing plays on the health and diversity of ecosystems and the welfare of the various species affected. Data from the fossil record of the Pleistocene - Holocene transition gives good insight into the role large herbivores play in ecosystem engineering and the possible effects of extinction (Cromsigt & Te Beest, 2014; Weigl & Knowles, 2014; Bakker et al., 2016). Furthermore, since most large herbivores a re grazers or mixed feeders (Shipley, 1999;Newman, 2000), an in - depth look at grasslands and related biomes, like steppes or savannas, clarifies the interactions of grazers and browsers with the other species of their environments. To highlight specific ec osystem services and the effects large herbivores have on their habitats, examples from the order Perissodactyla, the odd - toed ungulates, are considered. This order is highly relevant to the study of large herbivore extinction and diversity, since 13 of th e 16 species are endangered (with many extinct species represented in the fossil record), and they occur in ecosystems with a high level of biodiversity (Perissodactyl Preservation Fund, 2016). Megafauna in the Pleistocene The unique relationship between large herbivores and their habitats is not a new concept. The fossil evidence shows a long history of grazers and browsers maintaining a balance between the various plant species and the needs of the animals living off of th ose plants. Several studies have posited that large herbivores during the Pleistocene contributed to the vast amounts of open land during that period (Cromsigt & Te Beest, 2014). One such article suggests that the meadows in the Appalachian mountains were not cleared for human use in recent history but are artifacts left over from the time of the megafauna (Weigl & Knowles, 2014). The authors argue that the large herbivores that were abundant during the Pleistocene would have possessed the tolerance, mobili ty, strength, and diverse diets that permitted them to modify high - elevation habitats. They also claim that this theory of megafauna engineering would apply to the debate over Europe’s past. While some believe the continent was one unbroken forest, Weigl & Knowles (2014) cite studies supporting evidence that Europe was more a mosaic of grasslands, forests, and transition areas. Bakker et al. (2016) combined present day studies with paleo - data to determine the effects of megafauna on vegetation. The authors found several incidences that suggest large herbivores altered the landscape to maintain open, grass - dominated habitats. Wear patterns on teeth and tusks of mastodons ( Mammut americanum ) suggested bark - stripping behavior, which would have a marked effect o n woody plants. The reduction in tree cover and woody shrubs would then allow shade - intolerant plants, like grasses and forbs, to grow. Paleoecological evidence of geomorphological engineering by mammoths ( Mammuthus primigenius ), like digging, wallowing, o r trampling, would also have created open spaces for herbaceous vegetation. There is evidence for this theory in the pollen records of the Pleistocene - Holocene transition that show hardwoods increased immediately following the megafaunal decline. Johnson ( 2009) argues that the rise in water tables, loss of soil fertility, and increase in mosses in the mammoth steppes of North America and Europe during the Holocene was due to the loss of the megafauna. There is evidence that suggests the transition from step pe to tundra was due not just to climate change but to the lack of ecosystem engineering by the large herbivores that populated the biome. Grasses allow higher rates of transpiration of soil moisture than moss, and the cropping of the grass would have expo sed soil to rapid thawing and higher temperatures in warmer months. Without the grazing pressures, water tables rose, creating a favorable habitat for mosses, which insulate soil, keeping it cool and waterlogged. This, in turn, slowed the recycling of nutr ients, accumulating organic matter on the surface and reducing soil fertility. The author suggests that the engineering of the steppe by the megafauna may have damped the effects of climate warming on the vegetation, the effects of which were only truly fe lt after the disappearance of the engineers. Using evidence from the fossil record, Faith (2012) makes a good argument for present - day conservation practices that may provide Africa’s Cape mountain zebra ( Equus zebra zebra ) with better options in the futur e. With information gathered from historical data on megafauna in South Africa, the author makes a case for expansion
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