Megafauna

In zoology, megafauna (from Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower end being centered on humans, with other thresholds being more relative to the sizes of animals in an ecosystem, the spectrum of lower-end thresholds ranging from to . Large body size is generally associated with other traits, such as having a slow rate of reproduction and, in large herbivores, reduced or negligible adult mortality from being killed by predators.

Megafauna species have considerable effects on their local environment, including the suppression of the growth of woody vegetation and a consequent reduction in wildfire frequency. Megafauna also play a role in regulating and stabilizing the abundance of smaller animals.

During the Pleistocene, megafauna were diverse across the globe, with most continental ecosystems exhibiting similar or greater species richness in megafauna as compared to ecosystems in Africa today. During the Late Pleistocene, particularly from around 50,000 years ago onwards, most large mammal species became extinct, including 80% of all mammals greater than , while small animals were largely unaffected. This pronouncedly size-biased extinction is otherwise unprecedented in the geological record. Humans and climatic change have been implicated by most authors as the likely causes, though the relative importance of either factor has been the subject of significant controversy.

History

One of the earliest occurrences of the term "megafauna" is Alfred Russel Wallace's 1876 work ''The geographical distribution of animals''. He described the animals as "the hugest, and fiercest, and strangest forms". In the 20th and 21st centuries, the term usually refers to large animals. There are variations in thresholds used to define megafauna as a whole or certain groups of megafauna. Many scientific literature adopt Paul S. Martin's proposed threshold of to classify animals as megafauna. However, for freshwater species, is the preferred threshold. Some scientists define herbivorous terrestrial megafauna as having a weight exceeding , and terrestrial carnivorous megafauna as more than . Additionally, Owen-Smith coined the term megaherbivore to describe herbivores that weighed over , which has seen some use by other researchers.

Among living animals, the term megafauna is most commonly used for the largest extant terrestrial mammals, which include (but are not limited to) elephants, giraffes, hippopotamuses, rhinoceroses, and larger bovines. Of these five categories of large herbivores, only bovines are presently found outside of Africa and Asia, but all the others were formerly more wide-ranging, with their ranges and populations continually shrinking and decreasing over time. Wild equines are another example of megafauna, but their current ranges are largely restricted to the Old World, specifically in Africa and Asia. Megafaunal species may be categorized according to their dietary type: megaherbivores (e.g., elephants), megacarnivores (e.g., lions), and megaomnivores (e.g., bears).

Ecological strategy

Megafauna animals – in the sense of the largest mammals and birds – are generally ''K''-strategists, with high longevity, slow population growth rates, low mortality rates, and (at least for the largest) few or no natural predators capable of killing adults. These characteristics, although not exclusive to such megafauna, make them vulnerable to human overexploitation, in part because of their slow population recovery rates.

Evolution of large body size

One observation that has been made about the evolution of larger body size is that rapid rates of increase that are often seen over relatively short time intervals are not sustainable over much longer time periods. In an examination of mammal body mass changes over time, the maximum increase possible in a given time interval was found to scale with the interval length raised to the 0.25 power. This is thought to reflect the emergence, during a trend of increasing maximum body size, of a series of anatomical, physiological, environmental, genetic and other constraints that must be overcome by evolutionary innovations before further size increases are possible. A strikingly faster rate of change was found for large decreases in body mass, such as may be associated with the phenomenon of insular dwarfism. When normalized to generation length, the maximum rate of body mass decrease was found to be over 30 times greater than the maximum rate of body mass increase for a ten-fold change.

In terrestrial mammals

Subsequent to the Cretaceous–Paleogene extinction event that eliminated the non-avian dinosaurs about Ma (million years) ago, terrestrial mammals underwent a nearly exponential increase in body size as they diversified to occupy the ecological niches left vacant. Starting from just a few kg before the event, maximum size had reached ~ a few million years later, and ~ by the end of the Paleocene. This trend of increasing body mass appears to level off about 40 Ma ago (in the late Eocene), suggesting that physiological or ecological constraints had been reached, after an increase in body mass of over three orders of magnitude. However, when considered from the standpoint of rate of size increase per generation, the exponential increase is found to have continued until the appearance of '' Indricotherium'' 30 Ma ago. (Since generation time scales with ''body mass''^0.259, increasing generation times with increasing size cause the log mass vs. time plot to curve downward from a linear fit.)

Megaherbivores eventually attained a body mass of over . The largest of these, indricotheres and proboscids, have been hindgut fermenters, which are believed to have an advantage over foregut fermenters in terms of being able to accelerate gastrointestinal transit in order to accommodate very large food intakes. A similar trend emerges when rates of increase of maximum body mass per generation for different mammalian clades are compared (using rates averaged over macroevolutionary time scales). Among terrestrial mammals, the fastest rates of increase of ''body mass''^0.259 vs. time (in Ma) occurred in perissodactyls (a slope of 2.1), followed by rodents (1.2) and proboscids (1.1), all of which are hindgut fermenters. The rate of increase for artiodactyls (0.74) was about a third of the perissodactyls. The rate for carnivorans (0.65) was slightly lower yet, while primates, perhaps constrained by their arboreal habits, had the lowest rate (0.39) among the mammalian groups studied.

Terrestrial mammalian carnivores from several eutherian groups (the artiodactyl ''Andrewsarchus'' – formerly considered a mesonychid, the oxyaenid '' Sarkastodon'', and the carnivorans '' Amphicyon'' and ''Arctodus'') all reached a maximum size of about (the carnivoran ''Arctotherium'' and the hyaenodontid '' Simbakubwa'' may have been somewhat larger). The largest known metatherian carnivore, '' Proborhyaena gigantea'', apparently reached , also close to this limit. A similar theoretical maximum size for mammalian carnivores has been predicted based on the metabolic rate of mammals, the energetic cost of obtaining prey, and the maximum estimated rate coefficient of prey intake. It has also been suggested that maximum size for mammalian carnivores is constrained by the stress the humerus can withstand at top running speed.

Analysis of the variation of maximum body size over the last 40 Ma suggests that decreasing temperature and increasing continental land area are associated with increasing maximum body size. The former correlation would be consistent with Bergmann's rule, and might be related to the thermoregulatory advantage of large body mass in cool climates, better ability of larger organisms to cope with seasonality in food supply, or other factors; the latter correlation could be explained in terms of range and resource limitations. However, the two parameters are interrelated (due to sea level drops accompanying increased glaciation), making the driver of the trends in maximum size more difficult to identify.

In marine mammals

Since tetrapods (first reptiles, later mammals) returned to the sea in the Late Permian, they have dominated the top end of the marine body size range, due to the more efficient intake of oxygen possible using lungs. The ancestors of cetaceans are believed to have been the semiaquatic pakicetids, no larger than dogs, of about 53 million years (Ma) ago. By 40 Ma ago, cetaceans had attained a length of or more in ''Basilosaurus'', an elongated, serpentine whale that differed from modern whales in many respects and was not ancestral to them. Following this, the evolution of large body size in cetaceans appears to have come to a temporary halt and then to have backtracked, although the available fossil records are limited. However, in the period from 31 Ma ago (in the Oligocene) to the present, cetaceans underwent a significantly more rapid sustained increase in body mass (a rate of increase in ''body mass''^0.259 of a factor of 3.2 per million years) than achieved by any group of terrestrial mammals. This trend led to the largest animal of all time, the modern blue whale. Several reasons for the more rapid evolution of large body size in cetaceans are possible. Fewer biomechanical constraints on increases in body size may be associated with suspension in water as opposed to standing against the force of gravity, and with swimming movements as opposed to terrestrial locomotion. Also, the greater heat capacity and thermal conductivity of water compared to air may increase the thermoregulatory advantage of large body size in marine endotherms, although diminishing returns apply.

Among the toothed whales, maximum body size appears to be limited by food availability. Larger size, as in sperm and beaked whales, facilitates deeper diving to access relatively easily-caught, large cephalopod prey in a less competitive environment. Compared to odontocetes, the efficiency of baleen whales' filter feeding scales more favorably with increasing size when planktonic food is dense, making larger sizes more advantageous. The lunge feeding technique of rorquals appears to be more energy efficient than the ram feeding of balaenid whales; the latter technique is used with less dense and patchy plankton. The cooling trend in Earth's recent history may have generated more localities of high plankton abundance via wind-driven upwellings, facilitating the evolution of gigantic whales.

Cetaceans are not the only marine mammals to reach tremendous sizes. The largest mammal carnivorans of all time are marine pinnipeds, the largest of which is the southern elephant seal, which can reach more than in length and weigh up to . Other large pinnipeds include the northern elephant seal at , walrus at , and Steller sea lion at . The sirenians are another group of marine mammals which adapted to fully aquatic life around the same time as the cetaceans did. Sirenians are closely related to elephants. The largest sirenian was the Steller's sea cow, which reached up to in length and weighed , and was hunted to extinction in the 18th century.

In flightless birds

Because of the small initial size of all mammals following the extinction of the non-avian dinosaurs, nonmammalian vertebrates had a roughly ten-million-year-long window of opportunity (during the Paleocene) for evolution of gigantism without much competition. During this interval, apex predator niches were often occupied by reptiles, such as terrestrial crocodilians (e.g. '' Pristichampsus''), large snakes (e.g. '' Titanoboa'') or varanid lizards, or by flightless birds (e.g. '' Paleopsilopterus'' in South America). This is also the period when megafaunal flightless herbivorous gastornithid birds evolved in the Northern Hemisphere, while flightless paleognaths evolved to large size on Gondwanan land masses and Europe. Gastornithids and at least one lineage of flightless paleognath birds originated in Europe, both lineages dominating niches for large herbivores while mammals remained below (in contrast with other landmasses like North America and Asia, which saw the earlier evolution of larger mammals) and were the largest European tetrapods in the Paleocene.

Flightless paleognaths, termed ratites, have traditionally been viewed as representing a lineage separate from that of their small flighted relatives, the Neotropic tinamous. However, recent genetic studies have found that tinamous nest well within the ratite tree, and are the sister group of the extinct moa of New Zealand. Similarly, the small kiwi of New Zealand have been found to be the sister group of the extinct elephant birds of Madagascar. These findings indicate that flightlessness and gigantism arose independently multiple times among ratites via parallel evolution.

Predatory megafaunal flightless birds were often able to compete with mammals in the early Cenozoic. Later in the Cenozoic, however, they were displaced by advanced carnivorans and died out. In North America, the bathornithids '' Paracrax'' and '' Bathornis'' were apex predators but became extinct by the Early Miocene. In South America, the related phorusrhacids shared the dominant predatory niches with metatherian sparassodonts during most of the Cenozoic but declined and ultimately went extinct after eutherian predators arrived from North America (as part of the Great American Interchange) during the Pliocene