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Megaherbivore

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Large hippo next to a river
The common hippopotamus is an extant megaherbivore.

Megaherbivores (Greek μέγας megas "large" and Latin herbivora "herbivore"[1]) are large herbivores that can exceed 1,000 kg (2,200 lb) in weight. The earliest herbivores to reach such sizes like the pareiasaurian reptiles and dinocephalian therapsids appeared in the Permian period. During most of the Mesozoic, the megaherbivore niche was largely dominated by dinosaurs up until their extinction during the Cretaceous–Paleogene extinction event. After this period, small mammalian species evolved into large herbivores in the Paleogene. As part of the Late Pleistocene megafauna extinctions, 80% of megaherbivore species became extinct, with complete extinctions occurring in Europe, Australia and the Americas. Current megaherbivores include nine species of elephants, rhinos, hippos, and giraffes living in Africa and Asia.

Extant megaherbivores are keystone species in their environment. They defoliate the landscape and spread a greater number of seeds than other frugivores. Like most large mammals, extant megaherbivores are K-selected species and are characterized by their large size, relative immunity to predation, their effect on plant species, and their dietary tolerance.

Definition

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Megaherbivores are large herbivores that weigh more than a ton when fully grown.[2] They include both terrestrial and marine herbivores (i.e., manatees and dugongs),[1] and are classified as the largest type of (herbivorous) terrestrial megafauna.[3]

Evolution

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Permian

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The Early Permian caseid synapsids have been described as among the first megaherbivores.[4] Other major groups of Permian megaherbivores include dinocephalian therapsids (Middle Permian),[5] and pareiasaurian reptiles (Middle-Late Permian).[6] Megaherbivores became somewhat rare after the Permian–Triassic mass extinction event.[7][8]

Triassic

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Dicynodont therapsids were major megaherbivores form much of the Triassic period.[9] Lisowicia was the last dicynodont that lived and became extinct in the Late Triassic, and is the largest known dicynodont and non mammalian synapsid, being comparable in size to an Asian elephant.[7] Dinosaurs emerged as major megaherbivores during the later Triassic, cementing their dominance after the Triassic–Jurassic extinction event.[10]

Amniota
Synapsida

Dicynodontia (Permian, Triassic)

Mammals (small, until Paleogene)

Sauropsida

Pareiasauria (Permian)

Dinosauria (small, until Jurassic)

Jurassic

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Sauropodomorphs became the largest megaherbivores in the Jurassic. Other megaherbivorous groups included stegosaurs and ankylosaurs.[11]

Cretaceous

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From the Triassic to the Cretaceous, a diverse assemblage of megaherbivorous dinosaurs, such as sauropods,[12] occupied different ecological niches. Based on their dentition, ankylosaurs may have mainly consumed succulent plants, as opposed to nodosaurs, which were mainly browsers. It is thought that ceratopsids fed on rugged vegetation, due to their jaw being designed for a crushing effect. Studies on hadrosaur dentition concluded that they primarily fed on fruits.[13]

Paleogene

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Skeleton of Paraceratherium, a rhinocerotoid

Megaherbivore dinosaurs died out in the Cretaceous–Paleogene mass extinction.[14] For about 25 million years, the earth was void of large terrestrial herbivores that weighed more than 1 ton. After this period, small mammalian species evolved into large herbivores across every continent around 40 mya.[2] The largest of these animals were Paraceratheriidae and Proboscidea.[15] Other taxa included Brontotheriidae.[16] The Sirenia, aquatic megaherbivores, such as Dugongidae, Protosirenidae, and Prorastomidae were present in the Eocene.[17] Megaherbivores inhabited every major landmass in the Cenozoic and Pleistocene before the arrival of humans.[18]


Late Neogene

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During the late Miocene and Pliocene, African megaherbivore communities underwent a progressive, long-term decline. In eastern Africa, megaherbivore (mammals over 1000 kg) species richness began to fall around 4.6 million years ago, in the early Pliocene, more than a million years before the earliest evidence of hominin consumption of animal tissue (~3.3 Ma) and long before any sign of systematic hunting, ruling out an early anthropogenic origin for this decline.[19] Instead, it has been linked to environmental change, particularly the expansion of C4 grasslands and declining atmospheric CO2 concentrations, which favored grasses over woody vegetation.[19]

Body mass and abundance reconstructions based on fossil dental metrics confirm this pattern: African large-mammal communities older than ~4 Ma contained significantly more large-bodied individuals, with a greater share of total biomass concentrated in the largest size categories, than later communities. Over time, both individuals and biomass shifted toward smaller size categories, reflecting a gradual loss of the largest-bodied taxa. This process paralleled an approximately two-thirds decline in primary productivity since the late Miocene (>5 Ma), a globally observed trend that may have reduced the carrying capacity of large-mammal communities.[20]

More recent analyses of 23 million years of African ungulate diversification, based on neural network models, refine the ultimate cause of this diversity collapse. Starting around 7.2 Ma, both speciation and extinction rates accelerated broadly; speciation plateaued after 3.6 Ma, while extinction rose sharply at the onset of the Pleistocene (2.58 Ma), driving diversity losses across multiple herbivore guilds. However, extinction rates were intrinsically lower in larger-bodied lineages and in species with high-crowned (hypsodont) dentitions, contradicting the idea of extinction biased toward the largest species. The determining factor instead appears to be the intrinsically low speciation rate of megaherbivores themselves, further suppressed in some lineages by increasing aridification over the last 5 million years: the decline reflects insufficient speciation more than excess extinction.[21]


Pleistocene

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There were around 50 different species by the Late Pleistocene:[3]

Diprotodon, the largest marsupial to ever exist, was present across the entire Australian continent by the Late Pleistocene.[22] Elsewhere, megaherbivores like glyptodonts were grazing herbivores, that possessed no incisor or canine teeth, but had cheek teeth that would have been able to grind up tough vegetation. They inhabited habitats of South and North America.[23] Ground sloths were herbivores, with some being browsers,[24] others grazers,[25] and some intermediate between the two as mixed feeders.[26] Fossilized specimens were primarily found in South and North America, with one specimen being found as far north as Alaska.[27] Mammoths, like modern day elephants, had hypsodont molars. These features allowed mammoths to live an expansive life because of the availability of grasses and trees.[28] Today, nine of the 50 megaherbivore species persist. The Americas saw the worst decline in megaherbivores, with all 27 species going extinct.[3]

The Quaternary Extinction Event is an event where many species of megafauna (particularly mammals) went extinct. This event caused the disappearances of megaherbivores on most continents on Earth.[29] Climate change and the arrival of humans are considered likely causes of the extinctions.[30] It is thought that humans hunted megaherbivores to extinction, which then led to the extinction of the carnivores and scavengers which had preyed upon those animals.[31][32][33] Scientists have proposed that increasingly extreme weather—hotter summers and colder winters—referred to as "continentality", or related changes in rainfall caused the extinctions.[34]

Recent

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There are nine extant species of megaherbivores, found in Africa and Asia.[35][36] They include elephants, rhinos, hippos and giraffes.[35][37]: 1 [38] Elephants belong to the order Proboscidea; an order that has been around since the late Paleocene.[39] Hippopotamuses are the closest living relatives to cetaceans; soon after the common ancestor of whales and hippos diverged from even-toed ungulates, the lineages of cetaceans and hippopotamuses split apart.[40][41] Giraffidae are a sister taxon to Antilocapridae, with an estimated split of more than 20 million years ago, according to a 2019 genome study.[42] Rhinoceroses may originate from Hyrachyus, an animal whose remains date back to the late Eocene.[37]: 17 

Megaherbivores and other large herbivores are becoming less common throughout their natural distribution, impacting animal species within the ecosystem. This is mainly attributed to the destruction of their natural environment, agriculture, overhunting, and human invasion of their habitats.[43][44] As a consequence of their slow reproductive rate and the preference for targeting larger species, overexploitation poses the greatest threat to megaherbivores. As time progresses, it is thought that the situation will only worsen.[43]

Ecology of recent megaherbivory

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Browsers and grazers

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Living species exhibit the following adaptations: they have dietary tolerance, a strong effect on vegetation and with the exception of calves, face little threat from predators.[45][46][47]

Elephants and Indian rhinoceroses exhibit both grazing and browsing feeding habits. The hippopotamus and white rhinoceros prefer grazing herbivory, while giraffes and the three other rhinoceros species most often select browsing herbivory.[48] Mammalian megaherbivores predominantly consume graminoids. They prefer eating the leaves and stem of the plant, as well as its fruits. They also exhibit both foregut and hindgut fermentation, with rhinos, hippos, and elephants displaying the former and giraffes displaying the latter.[37]: 16  Their metabolic rate is lethargic, and as a result, digestion is slowed. During this prolonged digestion period, high-fiber plant matter is disintegrated.[38]

Due to their size, megaherbivores can defoliate the landscape; because of this, they are considered keystone species in their environment.[49] Megaherbivores affect the composition of plant species, which alters the movement and exchange of inorganic and organic matter back into the production of matter. They can open up areas through feeding behavior, which over time clears vegetation, including invasive alien plants. The number of seeds that megaherbivores spread is greater than that of other frugivores.[18][50] In addition, megaherbivore grazers, like the white rhino, have a profound impact on short grass. In one study, short grass became more infrequent after the elimination of white rhinos, which effected smaller grazers in the area.[48]

In a 2018 study, it was concluded that megaherbivores were not affected by the "landscape of fear," a landscape in which prey avoid certain hot-spot predation areas, thereby altering predator-frightened trophic cascades. Their feces were most apparent in closed, dense areas, indicating that they distribute resources to risky areas in this "landscape of fear".[51]

Interspecific interactions

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Most megaherbivore species are too big and powerful for most predators to kill.[38] Calves are, however, targeted by several predator species.[37]: 158  Giraffes are the most preyed upon megaherbivore, as it is not rare for lions to hunt adult giraffes in some places. The young are especially vulnerable, with a quarter to half of giraffe calves not reaching adulthood.[52][53] In Chobe National Park, lions have been recorded hunting young and sub-adult elephants.[54] Tigers are another known predator of young elephants.[55] Hippo calves may sometimes be prey items for lions, spotted hyenas and Nile crocodiles.[56]

Giraffes may flee or act in a non-aggressive manner, while white rhinos typically do not react to the presence of predators. Black and Indian rhinoceroses, elephants, and hippopotamuses on the other hand, react strongly to predators.[37]: 124–131 

Adaptations of extant megaherbivores

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Size

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Elephants are the largest members, weighing between 2.5 and 6.0 tons. Indian rhinos, white rhinos, and hippos usually weigh between 1.4 and 2.3 tons. The Javan and black rhino average 1–1.3 tons in weight. Giraffes are the smallest members, with a general weight range of 0.8–1.2 tons.[37]: 14 [57]

K-selection

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Extant megaherbivores are K-selected species, meaning they have high life expectancies, slow population growth, large offspring, lengthy pregnancies, and low mortality rates. They have selected slow reproduction to enhance their survival chances, and as a result, increase their lifespan.[58][59] Their large size offers protection from predators, but at the same, it decreases the degree at which they reproduce due to restricted food sources.[60][61] This slow population growth (elephants, for example, grow at a rate of 6–7%), indicates that populations may be drastically reduced if predation pressures are too great.[37]: 293  In stable environments, K-selection predominates as the ability to compete successfully for limited resources, and populations of K-selected organisms typically are very constant in number and close to the maximum that the environment can bear.[60][37]: 200 

Reproduction

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Giraffe calves do not stay with their mothers, they sit down and hide for most of the day, and their mothers briefly visit to feed them.[37]: 134 
Black rhino calves are vulnerable to predators, and stay close to their mothers for safety for 26 to 40 months.[37]: 136 

When females enter estrus, males will attempt to attract them as a mating partner. These breeding opportunities may be influenced by the hierarchical system of males. Giraffes and elephants mate for a relatively short time, while rhinos and hippos have a mating session lasting an extended period of time. Females have long gestation periods, between 8 and 22 months. Intervals between births vary between species, but the overall range is 1.3 to 4.5 years.[37]: 116–124 

They usually give birth to a single calf that is heavily reliant on females for food and protection. As they get older, the calf begins weaning while still suckling. When they reach juvenility, they are able to fend for themselves, but only to a certain extent. Females typically separate from their offspring by chasing them. Despite this, females may continue to interact with their progeny even after weaning.[37]: 133 

Lifespan and mortality

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Hippopotamuses and rhinoceroses can live to be 40 years old, while elephants can live longer than 60 years.[53] Giraffes have a lifespan of around 25 years.[37]: 158 

Around 2 to 5% of adult megaherbivores die each year. Males are more likely than females to die from wounds sustained during disputes. Occasionally, in times of drought, populations may significantly reduce, with calves being the most impacted during such times.[37]: 158 

See also

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References

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  1. 1 2 "Megaherbivore". Collins Dictionary. Retrieved 11 January 2024. Tropical seagrass meadows support a diversity of grazers spanning the meso-, macro-, and megaherbivore scales.
  2. 1 2 Onstein, Renske E.; Kissling, W. Daniel; Linder, H. Peter (2022-04-13). "The megaherbivore gap after the non-avian dinosaur extinctions modified trait evolution and diversification of tropical palms". Proceedings of the Royal Society B: Biological Sciences. 289 (1972) 20212633. doi:10.1098/rspb.2021.2633. ISSN 0962-8452. PMC 9006001. PMID 35414237.
  3. 1 2 3 Malhi, Yadvinder; Doughty, Christopher E.; Galetti, Mauro; Smith, Felisa A.; Svenning, Jens-Christian; Terborgh, John W. (2016-01-25). "Megafauna and ecosystem function from the Pleistocene to the Anthropocene". Proceedings of the National Academy of Sciences. 113 (4): 838–846. Bibcode:2016PNAS..113..838M. doi:10.1073/pnas.1502540113. ISSN 0027-8424. PMC 4743772. PMID 26811442.
  4. ↑ Werneburg, Ralf; Spindler, Frederik; Falconnet, Jocelyn; Steyer, Jean-Sebastien; Vianey-Liaud, Monique; Schneider, Jörg-W. (2022-07-18). "A New caseid Synapsid from the Permian (Guadalupian) of the Lodève basin (Occitanie, France)". Palaeovertebrata. 45 (2): e2. doi:10.18563/pv.45.2.e2.
  5. ↑ Kammerer, Christian F. (June 2011). "Systematics of the Anteosauria (Therapsida: Dinocephalia)". Journal of Systematic Palaeontology. 9 (2): 261–304. doi:10.1080/14772019.2010.492645. ISSN 1477-2019.
  6. ↑ Boitsova, Elizaveta A; Skutschas, Pavel P; Sennikov, Andrey G; Golubev, Valeriy K; Masuytin, Vladimir V; Masuytina, Olga A (2019-07-05). "Bone histology of two pareiasaurs from Russia (Deltavjatia rossica and Scutosaurus karpinskii) with implications for pareiasaurian palaeobiology". Biological Journal of the Linnean Society blz094. doi:10.1093/biolinnean/blz094. ISSN 0024-4066.
  7. 1 2 Sulej, Tomasz; Niedźwiedzki, Grzegorz (2019-01-04). "An elephant-sized Late Triassic synapsid with erect limbs". Science. 363 (6422): 78–80. Bibcode:2019Sci...363...78S. doi:10.1126/science.aal4853. ISSN 0036-8075. PMID 30467179.
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  9. ↑ Fiorelli, Lucas E.; Ezcurra, Martín D.; Hechenleitner, E. Martín; Argañaraz, Eloisa; Taborda, Jeremías R. A.; Trotteyn, M. Jimena; von Baczko, M. Belén; Desojo, Julia B. (2013-11-28). "The oldest known communal latrines provide evidence of gregarism in Triassic megaherbivores". Scientific Reports. 3 (1). doi:10.1038/srep03348. ISSN 2045-2322. PMC 3842779. PMID 24287957.
  10. ↑ Qvarnström, Martin; Vikberg Wernström, Joel; Wawrzyniak, Zuzanna; Barbacka, Maria; Pacyna, Grzegorz; Górecki, Artur; Ziaja, Jadwiga; Jarzynka, Agata; Owocki, Krzysztof; Sulej, Tomasz; Marynowski, Leszek; Pieńkowski, Grzegorz; Ahlberg, Per E.; Niedźwiedzki, Grzegorz (2024-12-12). "Digestive contents and food webs record the advent of dinosaur supremacy". Nature. 636 (8042): 397–403. doi:10.1038/s41586-024-08265-4. ISSN 0028-0836. PMC 11634772. PMID 39604731.
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  47. ↑ Ruggiero, Richard G. (March 1991). "Opportunistic predation on elephant calves". African Journal of Ecology. 29 (1): 86–89. Bibcode:1991AfJEc..29...86R. doi:10.1111/j.1365-2028.1991.tb00823.x. ISSN 0141-6707.
  48. 1 2 Hyvärinen, Olli (2022). Megaherbivores and Earth System Functioning: Landscape-scale Effects of White Rhino Loss on Vegetation, Fire and Soil Carbon Dynamics. Swedish University of Agricultural Sciences. p. 107. ISBN 978-91-7760-966-7.
  49. ↑ Gill, Jacquelyn L. (2013). "Ecological impacts of the late Quaternary megaherbivore extinctions". New Phytologist. 201 (4): 1163–1169. Bibcode:2014NewPh.201.1163G. doi:10.1111/nph.12576. PMID 24649488.
  50. ↑ Mungi, Ninad Avinash; Jhala, Yadvendradev V.; Qureshi, Qamar; le Roux, Elizabeth; Svenning, Jens-Christian (March 2023). "Megaherbivores provide biotic resistance against alien plant dominance". Nature Ecology & Evolution. 7 (10): 1645–1653. Bibcode:2023NatEE...7.1645M. doi:10.1038/s41559-023-02181-y. ISSN 2397-334X. PMID 37652995.
  51. ↑ le Roux, Elizabeth; Kerley, Graham I.H.; Cromsigt, Joris P.G.M. (August 2018). "Megaherbivores Modify Trophic Cascades Triggered by Fear of Predation in an African Savanna Ecosystem". Current Biology. 28 (15): 2493–2499.e3. Bibcode:2018CBio...28E2493L. doi:10.1016/j.cub.2018.05.088. ISSN 0960-9822. PMID 30033334.
  52. ↑ Lee, Derek E.; Bond, Monica L.; Kissui, Bernard M.; Kiwango, Yustina A.; Bolger, Douglas T. (2016-05-11). "Spatial variation in giraffe demography: a test of 2 paradigms". Journal of Mammalogy. 97 (4): 1015–1025. doi:10.1093/jmammal/gyw086. ISSN 1545-1542.
  53. 1 2 Bruton, Michael N. (2012-12-06). Alternative Life-History Styles of Animals. Springer Science & Business Media. p. 447. ISBN 978-94-009-2605-9.
  54. ↑ Power, R. John; Shem Compion, R.X. (April 2009). "Lion Predation on Elephants in the Savuti, Chobe National Park, Botswana". African Zoology. 44 (1): 36–44. doi:10.3377/004.044.0104. ISSN 1562-7020. S2CID 86371484.
  55. ↑ Thuppil, Vivek; Coss, Richard G. (2013-10-23). "Wild Asian elephants distinguish aggressive tiger and leopard growls according to perceived danger". Biology Letters. 9 (5) 20130518. doi:10.1098/rsbl.2013.0518. ISSN 1744-9561. PMC 3971691. PMID 24026347.
  56. ↑ Estes, R. (1992). The Behavior Guide to African Mammals: including hoofed mammals, carnivores, primates. University of California Press. pp. 222–226. ISBN 978-0-520-08085-0.
  57. ↑ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61 (3): 537–574. doi:10.4202/app.00136.2014. ISSN 0888-8892. S2CID 84294169.
  58. ↑ Manlik, Oliver (2019). "The Importance of Reproduction for the Conservation of Slow-Growing Animal Populations". Reproductive Sciences in Animal Conservation. Advances in Experimental Medicine and Biology. Vol. 1200. pp. 13–39. doi:10.1007/978-3-030-23633-5_2. ISBN 978-3-030-23632-8. ISSN 0065-2598. PMID 31471793. S2CID 201756810.
  59. ↑ Yuan, Rong; Hascup, Erin; Hascup, Kevin; Bartke, Andrzej (2023-11-01). "Relationships among Development, Growth, Body Size, Reproduction, Aging, and Longevity–Trade-Offs and Pace-Of-Life". Biochemistry (Moscow). 88 (11): 1692–1703. doi:10.1134/S0006297923110020. ISSN 1608-3040. PMC 10792675. PMID 38105191. S2CID 265507842.
  60. 1 2 Rafferty, John P. "K-selected species". Britannica. Retrieved 2 April 2017.
  61. ↑ Trimble, M. J.; Ferreira, S. M.; Van Aarde, R. J. (September 2009). "Drivers of megaherbivore demographic fluctuations: inference from elephants". Journal of Zoology. 279 (1): 18–26. doi:10.1111/j.1469-7998.2009.00560.x. ISSN 0952-8369.
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