Earth:Megaherbivore

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Short description: Megafauna subgroup


Large hippo next to a river
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. They first appeared 300 million years ago in the early Permian, in the form of synapsids. They were then replaced by megaherbivorous dinosaurs that went extinct in the Cretaceous-Paleogene extinction event. After this period, small mammalian species evolved into large herbivores in the Paleogene. During the Quaternary Extinction Event, megaherbivores disappeared on most continents on Earth. Recent megaherbivores include elephants, rhinos, hippos, and giraffes. There are nine extant species of terrestrial megaherbivores living in Africa and Asia. The African bush elephant is the largest extant species.

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

Definition

Megaherbivores are large herbivores that weigh more than 1 ton when fully grown.[2] They also include large marine herbivores.[1] They are a type of megafauna (>45 kg), and are the largest animals on land.[3]

Evolution

Permian

Dinodontosaurus skeleton

Megaherbivores first evolved in the early Permian (300 mya).[4] The earliest megaherbivores were synapsids; they became somewhat rare after the Permian–Triassic extinction event.[5][6] Taxa included dicynodonts such as Dinodontosaurus.[7] Pareiasaurs were other large herbivores present during this time.[8] The exact cause of the extinction remain unknown. It is thought that the main cause of extinction was the flood basalt volcanic eruptions that created the Siberian Traps,[9] which released sulfur dioxide and carbon dioxide, resulting in euxinia,[10] elevating global temperatures,[11] and acidifying the oceans.[12]

Triassic

Lisowicia was the last dicynodont that lived and became extinct in the Late Triassic.[5] Some scientists have proposed that there was never a Triassic–Jurassic extinction event, but others propose that the extinctions may have occurred earlier. Flood basalts are thought to be the primary driver of the extinctions, towards the end of the Triassic.[13] [14]

Amniota
Synapsida
 

Dicynodontia (Permian, Triassic) Dicynodont from PolandDB.jpg

 
 

Mammals (small, until Paleogene)

 
 
Sauropsida
 

Pareiasauria (Permian) Scutosaurus BW flipped.jpg

 
 

Dinosauria (small, until Jurassic)

 
 
 

Jurassic

The taxonomic structure then switched to sauropodomorphs. Other taxa included stegosaurs and ankylosaurs.[15] The change in taxonomy approximately occurred at the same time with the divergence of predominant vegetation and with extinctions. New taxa may have caused competitive exclusion (i.e. predominating and removing another taxa), or they may have adopted the ecological niche of extinct groups.[4][16]

Cretaceous

Dinosauria
Saurischia
 

(carnivorous dinosaurs)

 
 

Sauropoda AmpelosaurusDB.jpg

 
 
Ornithischia
Thyreophora
Ankylosauria
 

Nodosauridae Edmontonia dinosaur.png

 
 

Ankylosauridae Ankylosaurus magniventris reconstruction.png

 
 
Stegosauria

Stegosauridae Jiangjunosaurus junggarensis.png

 
 
Neornithischia
Ceratopsidae
 

Centrosaurinae Crittendenceratops shaded.jpg

 
 

Chasmosaurinae Utahceratops UDL.png

 
 
Hadrosauridae

Lambeosaurinae Tsintaosaurus spinorhinus NT.jpg

 
 
 
 

From the Triassic to the Cretaceous, a diverse assemblage of megaherbivorous dinosaurs, such as sauropods,[17] 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.[18]

Paleogene

Skeleton of a Paraceratheriid, a rhinocerotoid

Following the Cretaceous-Paleogene mass extinction, megaherbivore dinosaurs were extirpated from the face of the earth. One mechanism is thought to have played a major role: an extraterrestrial impact event in the Yucatán Peninsula.[19] For about 25 million years, the earth was void of large terrestrial herbivores that weighed more than 1 tonne. After this period, small mammalian species evolved into large herbivores. Herbivorous mammals had evolved to megaherbivore size across every continent around 40 mya.[2] The largest of these animals were Paraceratheriidae and Proboscidea.[20] Other taxa included Brontotheriidae.[21] The Sirenia, aquatic megaherbivores, such as Dugongidae, Protosirenidae, and Prorastomidae were present in the Eocene.[22] They inhabited every major landmass in the Cenozoic and Pleistocene before the arrival of humans.[4]

Pleistocene

Main page: Earth:Late Pleistocene extinctions

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

Mammalia
Marsupialia
 

Diprotodontidae (1 sp) Diprotodon optatum (2).jpg

 
 
Placentalia
Afrotheria
Sirenia

Manatees, Dugongs Dugong dugon Hardwicke white background.jpg

 
Proboscidea

Elephants, Mammoths (16 spp) 202003 Woolly mammoth.png

 
 
Boreoeutheria
Xenarthra
Cingulata

Giant glyptodonts (5 spp) Doedicurus BW.jpg

 
Pilosa

Giant sloths (8 spp) Thalassocnus carolomartini Restoration.png

 
 
Laurasiatheria
Panperissodactyla
 

Litopterna (1 sp) Macrauchenia (trunkless).jpg

 
 

Notoungulata toxodonts (1 sp) Toxodon.jpg

 
 

Perissodactyla Rhinocerotoids (9 spp) Rhino white background.jpg

 
 
Artiodactyla
 

Hippopotamuses (2 spp) Voyage en Abyssinie Plate 2 (white background).jpg

 
 
 

Giant camels (2 spp) Cladogram of Cetacea within Artiodactyla (Camelus bactrianus).png

 
 

Giant bovids (3 spp) Birds and nature (1901) (14562088237) white background.jpg

 
 
 
 
 
 
 

Diprotodon was present across the entire Australian continent by the Late Pleistocene.[23] Glyptodonts were grazing herbivores. Like many other xenarthrans, they had no incisor or canine teeth, but had cheek teeth that would have been able to grind up tough vegetation. [24] Ground sloths were herbivores, with some being browsers,[25] others grazers,[26] and some intermediate between the two as mixed feeders.[27] Mammoths, like modern day elephants, have hypsodont molars. These features also allowed mammoths to live an expansive life because of the availability of grasses and trees.[28]

Today, nine of the 50 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 disappearance of megaherbivores on most continents on Earth.[29] Climate change and the arrival of humans could be the 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

Placentalia
Afrotheria
 
Sirenia

Manatees, Dugongs (4 spp) Dugong dugon Hardwicke white background.jpg

 
Proboscidea

Elephants (3 spp) Elephant white background.png

 
 
 
Boreoeutheria
 
Perissodactyla

Rhinoceroses (4 spp) Rhino white background.jpg

 
Artiodactyla
 

Giraffes (1 sp) Giraffa camelopardalis Brockhaus white background.jpg

 
 

Hippopotamuses (1 sp) Voyage en Abyssinie Plate 2 (white background).jpg

 
 
 
 
 

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, 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 dates back to the late Eocene.[37](p17)

Megaherbivores and other large herbivores are becoming less common throughout their natural distribution, which is having an impact on 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 grow worse.[43]

Ecology of recent megaherbivory

Browsers and grazers

Living species exhibit the following adaptations: they have dietary tolerance, have a strong effect on vegetation and with the exception of calves, they 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](p16)

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. The number of seeds that megaherbivores spread is greater than that of other frugivores.[4] 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, affecting smaller grazers in the process.[48] Their metabolic rate is lethargic, and as a result, digestion is slowed. During this prolonged digestion period, high-fiber plant matter is disintegrated.[38]

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".[50]

Interspecific interactions

Most megaherbivore species are too big and powerful for most predators to kill.[38] Calves are, however, targeted by several predator species.[37](p158) Giraffes are susceptible to predation, and it's not rare for lions and spotted hyenas to hunt adult giraffes in some places. The young are especially vulnerable, with a quarter to half of giraffe calves not reaching adulthood.[51][52] In Chobe National Park, lions have been recorded hunting young and sub-adult elephants.[53] Tigers are another known predator of young elephants.[54] Hippo calves may sometimes be prey items for lions, spotted hyenas and Nile crocodiles.[55]

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 react strongly to predators.[37](pp124-131)

Adaptations of extant megaherbivores

Size

Elephants are the largest members, weighing between 2.5–6.0 tons. Indian rhinos, white rhinos and hippos usually weigh between 1.4–2.3 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](p14)[56]

K-selection

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.[57][58] Their large size offers protection from predators, but at the same, it decreases the degree at which they reproduce due to restricted food sources.[59][60] Due to their slow population growth (elephants, for example, grow at a rate of 6–7%), populations may be drastically reduced if the population growth rate is not greater than the rate of predation.[37](p293) 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.[59][37](p200) Due to the effects of human activity, population densities are currently sparse.[37](p21)

Reproduction

Giraffe calves don't stay with their mothers, they sit down and hide for most of the day, and their mothers briefly visit to feed them.[37](p134)
Black rhino calves are vulnerable to predators, and stay close to their mothers for safety for 26 to 40 months.[37](p136)

When females enter estrus, males will attempt to attract and mate with them. Breeding opportunities may be influenced by the hierarchical system within 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](p116-124)

They usually give birth to a single calf. Calves are born helpless and heavily rely 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](p133)

Lifespan and mortality

Hippopotamuses and rhinoceroses can live to be 40 years old, while elephants can live longer than 60 years.[52] Giraffes have a lifespan of around 25 years.[37](p158)

Around 2 to 5% of adult megaherbivores die each year. Males are more likely than females to die from wounds sustained during disputes. Giraffes are the most preyed-upon megaherbivore species. Occasionally, in times of drought, populations may significantly reduce, with calves being the most impacted.[37](p158)

See also

  • Australian megafauna
  • Deep-sea gigantism
  • Largest and heaviest animals
  • List of extant megaherbivores

References

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  2. ↑ 2.0 2.1 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). doi:10.1098/rspb.2021.2633. ISSN 0962-8452. PMID 35414237. PMC 9006001. http://dx.doi.org/10.1098/rspb.2021.2633. 
  3. ↑ 3.0 3.1 3.2 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. doi:10.1073/pnas.1502540113. ISSN 0027-8424. PMID 26811442. Bibcode: 2016PNAS..113..838M. 
  4. ↑ 4.0 4.1 4.2 4.3 Bocherens, Hervé (2018). "The Rise of the Anthroposphere since 50,000 Years: An Ecological Replacement of Megaherbivores by Humans in Terrestrial Ecosystems?". Frontiers in Ecology and Evolution 6. doi:10.3389/fevo.2018.00003. ISSN 2296-701X. 
  5. ↑ 5.0 5.1 Sulej, Tomasz; Niedźwiedzki, Grzegorz (2019-01-04). "An elephant-sized Late Triassic synapsid with erect limbs". Science 363 (6422): 78–80. doi:10.1126/science.aal4853. ISSN 0036-8075. PMID 30467179. Bibcode: 2019Sci...363...78S. http://dx.doi.org/10.1126/science.aal4853. 
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  26. ↑ van Geel, Bas; van Leeuwen, Jacqueline F.N.; Nooren, Kees; Mol, Dick; den Ouden, Natasja; van der Knaap, Pim W.O.; Seersholm, Frederik V.; Rey-Iglesia, Alba et al. (January 2022). "Diet and environment of Mylodon darwinii based on pollen of a Late-Glacial coprolite from the Mylodon Cave in southern Chile" (in en). Review of Palaeobotany and Palynology 296: 104549. doi:10.1016/j.revpalbo.2021.104549. Bibcode: 2022RPaPa.29604549V. 
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  33. ↑ Martin, P. S. (1989). "Prehistoric overkill: A global model". in Martin, P.S.. Quaternary extinctions: A prehistoric revolution. Tucson, AZ: University of Arizona Press. pp. 354–404. ISBN 978-0-8165-1100-6. 
  34. ↑ Birks, H.H. (1973). "Modern macrofossil assemblages in lake sediments in Minnesota". in Birks, H.J.B.. Oxford: Blackwell Scientific. ISBN 0-632-09120-7. 
  35. ↑ 35.0 35.1 (in en) 'Wildlife' Review. U.S. Department of the Interior, Fish and Wildlife Service. 1989. https://books.google.com/books?id=XcJGAQAAMAAJ&dq=Megaherbivores&pg=PA524. 
  36. ↑ Wells, Harry B. M.; Crego, Ramiro D.; Opedal, Øystein H.; Khasoha, Leo M.; Alston, Jesse M.; Reed, Courtney G.; Weiner, Sarah; Kurukura, Samson et al. (2021). "Experimental evidence that effects of megaherbivores on mesoherbivore space use are influenced by species' traits" (in en). Journal of Animal Ecology 90 (11): 2510–2522. doi:10.1111/1365-2656.13565. ISSN 0021-8790. PMID 34192343. Bibcode: 2021JAnEc..90.2510W. 
  37. ↑ 37.00 37.01 37.02 37.03 37.04 37.05 37.06 37.07 37.08 37.09 37.10 37.11 37.12 37.13 37.14 Owen-Smith, R. Norman (1988) (in en). Megaherbivores: The Influence of Very Large Body Size on Ecology. Cambridge University Press. ISBN 978-0-521-42637-4. https://books.google.com/books?id=1jT72JdLVIcC&q=megaherbivores. 
  38. ↑ 38.0 38.1 38.2 Ray, Justina; Redford, Kent H.; Steneck, Robert; Berger, Joel (2013-04-09) (in en). Large Carnivores and the Conservation of Biodiversity. Island Press. pp. 87. ISBN 978-1-59726-609-3. https://books.google.com/books?id=ndb0QOvq2LYC&dq=megaherbivores&pg=PA87. 
  39. ↑ Gheerbrant, E. (2009). "Paleocene emergence of elephant relatives and the rapid radiation of African ungulates". Proceedings of the National Academy of Sciences of the United States of America 106 (26): 10717–10721. doi:10.1073/pnas.0900251106. PMID 19549873. Bibcode: 2009PNAS..10610717G. 
  40. ↑ Gatesy, J. (1 May 1997). "More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen". Molecular Biology and Evolution 14 (5): 537–543. doi:10.1093/oxfordjournals.molbev.a025790. PMID 9159931. 
  41. ↑ Boisserie, Jean-Renaud; Lihoreau, Fabrice; Brunet, Michel (2005). "The position of Hippopotamidae within Cetartiodactyla". Proceedings of the National Academy of Sciences 102 (5): 1537–1541. doi:10.1073/pnas.0409518102. PMID 15677331. Bibcode: 2005PNAS..102.1537B. 
  42. ↑ Chen, Lei; Qiu, Qiang; Jiang, Yu; Wang, Kun; Lin, Zeshan et al. (2019-06-21). "Large-scale ruminant genome sequencing provides insights into their evolution and distinct traits" (in en). Science 364 (6446). doi:10.1126/science.aav6202. ISSN 0036-8075. PMID 31221828. Bibcode: 2019Sci...364.6202C. https://www.science.org/doi/10.1126/science.aav6202. 
  43. ↑ 43.0 43.1 Ripple, William J.; Newsome, Thomas M.; Wolf, Christopher; Dirzo, Rodolfo; Everatt, Kristoffer T. et al. (May 2015). "Collapse of the world's largest herbivores". Science Advances 1 (4): e1400103. doi:10.1126/sciadv.1400103. ISSN 2375-2548. PMID 26601172. Bibcode: 2015SciA....1E0103R. 
  44. ↑ Schowanek, Simon D.; Davis, Matt; Lundgren, Erick J.; Middleton, Owen; Rowan, John; Pedersen, Rasmus Ø.; Ramp, Daniel; Sandom, Christopher J. et al. (April 2021). Lyons, Kathleen. ed. "Reintroducing extirpated herbivores could partially reverse the late Quaternary decline of large and grazing species" (in en). Global Ecology and Biogeography 30 (4): 896–908. doi:10.1111/geb.13264. ISSN 1466-822X. Bibcode: 2021GloEB..30..896S. https://onlinelibrary.wiley.com/doi/10.1111/geb.13264. 
  45. ↑ Owen-Smith, N. (2013). "Megaherbivores". Encyclopedia of Biodiversity (Second ed.). pp. 223–229. doi:10.1016/B978-0-12-384719-5.00358-0. ISBN 9780123847201. https://www.researchgate.net/publication/323825899. 
  46. ↑ Clauss, Marcus; Frey, R; Kiefer, B; Lechner-Doll, M. (2003). "The maximum attainable body size of herbivorous mammals: Morphophysiological constraints on foregut, and adaptations of hindgut fermenters". Oecologia 136 (1): 14–27. doi:10.1007/s00442-003-1254-z. PMID 12712314. Bibcode: 2003Oecol.136...14C. https://www.researchgate.net/publication/7448963. 
  47. ↑ Ruggiero, Richard G. (March 1991). "Opportunistic predation on elephant calves" (in en). African Journal of Ecology 29 (1): 86–89. doi:10.1111/j.1365-2028.1991.tb00823.x. ISSN 0141-6707. Bibcode: 1991AfJEc..29...86R. https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2028.1991.tb00823.x. 
  48. ↑ 48.0 48.1 Hyvärinen, Olli (2022) (in en). Megaherbivores and Earth System Functioning: Landscape-scale Effects of White Rhino Loss on Vegetation, Fire and Soil Carbon Dynamics. Swedish University of Agricultural Sciences. pp. 107. ISBN 978-91-7760-966-7. https://books.google.com/books?id=7w9fzwEACAAJ. 
  49. ↑ Gill, Jacquelyn L. (2013). "Ecological impacts of the late Quaternary megaherbivore extinctions". New Phytologist 201 (4): 1163–1169. doi:10.1111/nph.12576. PMID 24649488. https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.12576#:~:text=The%20surviving%20megaherbivores%20(see%20Box,ecosystems%20(Johnson%2C%202009). 
  50. ↑ 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. doi:10.1016/j.cub.2018.05.088. ISSN 0960-9822. PMID 30033334. http://dx.doi.org/10.1016/j.cub.2018.05.088. 
  51. ↑ 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. http://dx.doi.org/10.1093/jmammal/gyw086. 
  52. ↑ 52.0 52.1 Bruton, Michael N. (2012-12-06) (in en). Alternative Life-History Styles of Animals. Springer Science & Business Media. pp. 447. ISBN 978-94-009-2605-9. https://books.google.com/books?id=6Er1CAAAQBAJ&dq=Megaherbivores&pg=PA440. 
  53. ↑ 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. http://dx.doi.org/10.3377/004.044.0104. 
  54. ↑ Thuppil, Vivek; Coss, Richard G. (2013-10-23). "Wild Asian elephants distinguish aggressive tiger and leopard growls according to perceived danger" (in en). Biology Letters 9 (5): 20130518. doi:10.1098/rsbl.2013.0518. ISSN 1744-9561. PMID 24026347. 
  55. ↑ 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. https://archive.org/details/isbn_0520080858/page/222. 
  56. ↑ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans". Acta Palaeontologica Polonica 61 (3): 537–574. doi:10.4202/app.00136.2014. ISSN 0888-8892. https://www.app.pan.pl/archive/published/app61/app001362014.pdf. 
  57. ↑ 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. 1200. pp. 13–39. doi:10.1007/978-3-030-23633-5_2. ISBN 978-3-030-23632-8. https://pubmed.ncbi.nlm.nih.gov/31471793/. 
  58. ↑ 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" (in en). Biochemistry (Moscow) 88 (11): 1692–1703. doi:10.1134/S0006297923110020. ISSN 1608-3040. PMID 38105191. PMC 10792675. https://doi.org/10.1134/S0006297923110020. 
  59. ↑ 59.0 59.1 Rafferty, John P.. "K-selected species". Britannica. https://www.britannica.com/science/K-selected-species. Retrieved 2 April 2017. 
  60. ↑ Trimble, M. J.; Ferreira, S. M.; Van Aarde, R. J. (September 2009). "Drivers of megaherbivore demographic fluctuations: inference from elephants" (in en). Journal of Zoology 279 (1): 18–26. doi:10.1111/j.1469-7998.2009.00560.x. ISSN 0952-8369. https://zslpublications.onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.2009.00560.x. 



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