Everything about The Great American Interchange totally explained
The
Great American Interchange was an important
paleozoogeographic event in which land and freshwater
fauna migrated from
North America via
Central America to
South America and vice versa, as the volcanic
Isthmus of Panama rose up from the sea floor and bridged the
continents. The migration peaked dramatically around 3 million years (
Ma) ago (in the
Piacenzian, the first half of the Upper
Pliocene).
It resulted in the joining of the
Neotropic (roughly South America) and
Nearctic (roughly North America) definitively to form the
Americas. The interchange is visible from observation of both
stratigraphy and nature (
neontology). Its most dramatic effect is on the
zoogeography of
mammals but it also gave an opportunity for non-flying
birds,
arthropods,
reptiles,
amphibians and even freshwater
fish to migrate.
South America's endemic fauna
After the late
Mesozoic breakup of
Gondwana, South America spent most of the
Cenozoic era as an island continent whose "splendid isolation" allowed its fauna to evolve into forms found nowhere else on earth. Its
endemic mammals initially consisted of
marsupials,
xenarthrans (for example,
armadillos,
anteaters and
sloths, like the giant ground sloth
Megatherium), and a diverse group of
native ungulates:
notoungulates (the "southern
ungulates"),
litopterns,
astrapotheres (for example
Trigonostylops,
Astrapotherium), and
pyrotheres (for example
Pyrotherium).
Marsupials may have traveled (via
Gondwanan land connections) from South America through
Antarctica to
Australia and/or vice versa in the late
Cretaceous or early
Tertiary. One living S. American marsupial, the
Monito del Monte, is believed to be more closely related to
Australian marsupials than to
other South American marsupials. A
61-Ma-old platypus-like monotreme fossil from
Patagonia may represent another Australian immigrant. It appears that
ratites (relatives of S. American
tinamous) migrated by this route around the same time, more likely in the direction from S. America towards Australia/
New Zealand. Other taxa that may have dispersed by the same route (if not by flying or floating) are
parrots,
chelid turtles and (extinct)
meiolaniid turtles.
The marsupials present in South America included didelphimorphs (
opossums and relatives), but many
larger predatory forms also existed, like the
borhyaenids and the
sabertooth Thylacosmilus. The marsupials shared the
ecological niches for large predators with fearsome flightless "terror birds" (
phorusrhacids), whose closest
extant relatives are the
seriemas.
The notoungulates and litopterns had many strange forms, like
Macrauchenia, a camel-like litoptern with a small
proboscis. They also produced a number of familiar-looking body types that represent examples of
parallel or
convergent evolution: one-toed
Thoatherium had legs like those of a horse,
Pachyrukhos resembled a rabbit,
Homalodotherium was a semi-bipedal clawed browser like a
chalicothere, and horned
Trigodon looked like a
rhino. Both groups started evolving in the Lower
Paleocene, possibly from
condylarth stock, diversified, dwindled before the great interchange, and went
extinct in the
Pleistocene. The pyrotheres and astrapotheres were also strange but were less diverse and disappeared earlier, well before the interchange.
The North American fauna was a pretty typical
boreoeutherian one (supplemented with
Afrotherian
proboscideans).
Island-hopping ‘waif dispersers’
The invasions of South America started at least 31.5 Ma ago (late
Eocene/early
Oligocene), when
cavimorph rodents arrived. This gave rise to – among others –
capybaras,
chinchillas,
viscachas, and
New World porcupines. (The independent development of
spines by New and
Old World porcupines is another example of parallel evolution.) This invasion most likely came from Africa. The crossing from
West Africa to the northeast corner of
Brazil was much shorter then due to
continental drift, and may have been aided by
island-hopping (for example via
St. Paul's Rocks, if they were an inhabitable island at the time) and westward oceanic currents. Crossings of the ocean were accomplished when at least one fertilised female (more commonly a group of animals) accidentally floated over on
driftwood or
mangrove rafts. (
Island-hopping cavimorphs would subsequently colonize the
West Indies as far as the
Bahamas).
A little later (at least 25 Ma ago)
primates followed. The ancestor of the South American monkeys is believed to have arrived from Africa in a fashion similar to the rodents.
The primates capable of migrating had to be small. These gave rise to the
New World monkeys (Platyrrhini). (Not long after arriving, monkeys apparently most closely related to
titis island-hopped to
Cuba,
Hispaniola
and
Jamaica.) The South American cavimorph rodents and monkeys are both believed to be
clades (for example,
monophyletic).
Tortoises also arrived in South America in the Oligocene. It was long thought that they'd come from N. America, but a recent comparative genetic analysis concludes that S. American members of
Geochelone are actually most closely related to African
hingeback tortoises. Tortoises are aided in oceanic dispersal by their ability to float with their heads up, and to survive up to six months without food or water.
The earliest mammalian arrivals from North America were
carnivorous procyonids that island-hopped from Central America prior to the formation of a land bridge, around 7 Ma ago. Some South American procyonids then diversified into forms now extinct (for example the "dog-coati"
Cyonasua, which evolved into the
bear-like
Chapalmalania). However, all extant procyonid genera appear to have originated in North America. The procyonids were followed to S. America by island-hopping
sigmodontine rodents,
peccaries and
hog-nosed skunks.
Similarly, mylodontid and megalonychid
ground sloths island-hopped to North America by 9 Ma ago., and may have reached N. America by that route, since the oldest N. American fossils are from
Florida.
The Great American Biotic Interchange
The formation of the Isthmus of Panama led to the last and most conspicuous wave, the great interchange, around 3 Ma ago. This included the immigration of North American ungulates (including
llamas,
tapirs and
horses),
proboscideans (such as
mastodons),
carnivorans (including
felids like
cougars and
saber-toothed cats,
canids, and
bears) and a number of types of
rodents
into South America.
In general, the initial net migration was symmetrical. Later on, however, the Neotropic species proved far less successful than the Nearctic. This "bad luck" happened both ways. Northwardly migrating animals often were not able to compete for resources as well as the North American species already occupying the same ecological niches; even when they became established, they usually didn't diversify much. Southwardly migrating Nearctic species diversified more, and are thought to have caused the
extinction of much of the South American fauna. (There were no extinctions in N. America obviously attributable to S. American immigrants.) Although terror birds were initially able to invade N. America, this success was temporary; all of the large Neotropic avian and marsupial predators ultimately disappeared. South America's
native ungulates also fared very poorly, with only several of the largest forms,
Macrauchenia and a few
toxodonts, withstanding the northern onslaught. (Among the notoungulates, the
mesotheriids and
hegetotheriids did manage to survive into the Pleistocene.) Its
small marsupials fared better, while the
primitive-looking
xenarthrans proved to be surprisingly competitive. The African immigrants, the cavimorph rodents and platyrrhine monkeys, generally held their own during the interchange, although the largest rodents (for example the
dinomyids) seem to have disappeared. With the exception of the
North American porcupine and several extinct porcupines and capybaras, however, they didn't migrate past
Central America.
The presence of armadillos, opossums, and porcupines in the
United States today is explained by the Great American Interchange. Opossums and porcupines were among most successful northward migrants, reaching as far as
Canada. While only one example each of xenarthrans, marsupials and cavimorph rodents currently lives in North America, 32 species from these
taxa are
present in tropical Central America. Prior to the end-
Pleistocene extinctions, most major groups of xenarthrans were established in Central or North America. Among the
megafauna, ground sloths were notably successful invaders of North America;
Megalonyx spread as far north as
Alaska. Generally speaking, however, the dispersal and subsequent explosive
adaptive radiation of
sigmodontine rodents through South America was much more successful (both spatially and by number of species) than any northward migration of South American mammals.
Reasons for success or failure
The eventual triumph of the Nearctic migrants was ultimately based on
geography, which played into the hands of the northern invaders in two crucial respects. The first was a matter of
climate. Obviously, any species that reached
Panama from either direction had to be able to tolerate . Those migrating southward would then be able to occupy much of South America without encountering climates that were markedly different. However, northward migrants would begin to encounter drier and/or cooler conditions as soon as they reached the doorstep of North America. The challenge this climatic asymmetry (see map) presented was particularly acute for Neotropic species specialized for
tropical rainforest environments, who had little prospect of penetrating beyond Central America.
The second and more important advantage geography gave to the northerners is related to the land area available for their ancestors to evolve in. During the Cenozoic, North America was periodically connected to
Eurasia via
Beringia, allowing multiple migrations back and forth to unite the faunas of the two continents. Eurasia was
connected in turn to Africa, which contributed further to the species that made their way to North America. South America, on the other hand, was connected to Antarctica and Australia, two much smaller continents, only in the earliest part of the Cenozoic, and this land connection doesn't seem to have carried much traffic (apparently no mammals other than marsupials and perhaps a few monotremes ever migrated by this route). Effectively, this means that northern hemisphere species arose over a land area roughly six times larger than was available to S. American species. This calculation may not be entirely fair, in that migrations between continents would have been more difficult and less frequent than migrations within S. America. Nevertheless, it's clear that N. American species were products of a larger and more competitive arena, where
evolution would have proceeded more rapidly. They tended to be more efficient and
brainier, generally able to outrun and outwit their S. American counterparts. These advantages can be clearly seen in the cases of ungulates and their predators, where S. American forms were replaced wholesale by the invaders.
Against this backdrop, the ability of S. America's
xenarthrans to compete effectively against the northerners represents a special case. The explanation for the xenarthrans' success lies in their idiosyncratic approach to defending against predation, based on possession of
body armor and/or formidable
claws. The xenarthrans didn't need to be fleet-footed or quick-witted to survive. Such a strategy may have been forced on them by their low
metabolic rate (the lowest among the
therians). Their low metabolic rate may in turn have been advantageous in allowing them to specialize on less abundant and/or less nutritious food sources.
End-Pleistocene extinctions
At the end of the
Pleistocene epoch, about 12,000 years ago, three dramatic developments occurred in the Americas at roughly the same time (geologically speaking).
Paleoindians invaded and occupied the
New World, the
last glacial period came to an end, and a large fraction of the
megafauna of both
North and
South America went extinct. This
wave of extinctions swept off the face of the
Earth many of the successful participants of the Great American Interchange, as well as other species that hadn't migrated. All the proboscideans, equids, lions,
Smilodon species, dire wolves, ground sloths, glyptodonts, and pampatheres of both continents disappeared. The last of the South and Central American notoungulates and litopterns died out, as well as North America's
native cheetahs,
scimitar cats,
giant beavers and many of its
bovid,
cervid,
tayassuid and
antilocaprid ungulates. Some groups survived in their adopted homes but disappeared over most or all of their original range, for example South American camelids, tapirs and tremarctine bears (cougars and jaguars may have been temporarily reduced to S. American ranges also). Others, such as capybaras, survived in their original range but died out in areas they'd migrated to.
The near simultaneity of the megafaunal extinctions with the glacial retreat and the
peopling of the Americas has led to proposals that both climate change and human hunting played a role. However, a number of considerations suggest that human activities were pivotal. The extinctions didn't occur selectively in the climatic zones that would have been most affected by the warming trend. The climate change took place worldwide, but had little effect on the megafauna in areas like
Africa and
South Asia, where megafaunal species had coevolved with
humans. Numerous
very similar
glacial retreats
had occurred previously within the
ice age of the last several Ma without ever producing comparable waves of extinction in the Americas or anywhere else. Similar megafaunal extinctions have occurred on other recently populated land masses (for example
Australia,
Madagascar, New Zealand, and many smaller islands around the world, such as
Cyprus) at different times that correspond closely to the first arrival of humans at each location. Additionally, on sizable islands far enough offshore from newly occupied territory to escape immediate human colonization, megafaunal species sometimes survived for thousands of years after they became extinct on the mainland; examples include
wooly mammoths on
Wrangel Island and
Saint Paul Island,
ground sloths on the
Antilles,
Steller's sea cows off the
Commander Islands, and
meiolaniid turtles on
Lord Howe Island and
New Caledonia. The glacial retreat may have played a primarily indirect role in the extinctions by simply facilitating the movement of humans southeastward from
Beringia down to N. America.
South American species that migrated to North America
Extant or
extinct North American
taxa whose ancestors migrated out of South America during the last 10
Ma:
South American species that migrated only as far as Central America
Extant or extinct Central American taxa whose ancestors migrated out of South America during the last 10
Ma of genera
Caecilia,
Dermophis,
Gymnopis and
Oscaecilia
Dendrobatid Poison Dart Frogs
Boas (snakes of the subfamily Boinae)
other Opossums (12 additional extant species, listed on )
Northern Naked-tailed Armadillo (Cabassous centralis)
Hoffmann's Two-toed Sloth (Choloepus hoffmanni)
Three-toed Sloths (Bradypus variegatus and B. pygmaeus)
Silky and Giant Anteaters, and Northern Tamandua
Toxodonts (†Mixotoxodon, a rhino-like notoungulate)
Rothschild's and Mexican Hairy Dwarf Porcupines (Coendou rothschildi and Sphiggurus mexicanus)
other Caviomorph Rodents (9 additional extant species, listed on )
Platyrrhine Monkeys (up to 10 extant species, listed on )
other Vampire Bats (all 3 extant species)
Toucans
Tinamous
North American species that migrated to South America
Extant or extinct South American taxa whose ancestors migrated out of North or Central America during the last 10 Ma and Oedipina
Coral Snakes
Rattlesnakes
other Pit Vipers (for example, ancestors of bushmasters and lanceheads)
Small-eared Shrews (genus Cryptotis; only present in NW S. America - Colombia, Venezuela, Ecuador, Peru)
Geomyid Pocket Gophers (one species, Orthogeomys thaeleri, in Colombia)
Heteromyid Mice (genus Heteromys; only present in NW S. America - Colombia, Venezuela, Ecuador])
Cricetid (primarily Sigmodontine) Rats and Mice
Tree Squirrels of genera Sciurus, Microsciurus, and Sciurillus
Cottontail Rabbits (Sylvilagus brasiliensis, S. floridanus and S. varynaensis)
Tapirs
Equids (†Plesippus, †Hippidion)
Peccaries
Deer of genera Odocoileus, Blastocerus, Ozotoceros, Mazama, Pudu, and Hippocamelus
Camelids
American Mastodon (†Mammut americanum)
Gomphotheres (†Cuvieronius, elephant relatives)
Otters of genera Lontra and Pteronura
other Mustelids, from subfamily Mustelinae, of genera Eira, Galictis, Lyncodon and Mustela
Hog-Nosed Skunks (Conepatus chinga, C. humboldtii and C. semistriatus)
Procyonids (raccoons, coatis, kinkajous and olingos)
Spectacled Bear (Tremarctos ornatus)
other Short-Faced Bears of subfamily Tremarctinae (†Arctotherium brasilense, †Arctotherium latidens)
Wolves (for example †Canis dirus, †C. ambrusteri)
Gray Fox (Urocyon cinereoargenteus, only present in NW S. America - Colombia, Venezuela)
other Canids (genera Atelocynus, Cerdocyon, Pseudalopex, Chrysocyon, and Speothos)
Cougar (Puma concolor) and jaguarundi (P. yaguarondi)
Jaguar (Panthera onca)
Saber-Toothed Cats (†Smilodon gracilis, †S. fatalis, †S. populator)
American Lion (†Panthera leo atrox)
all members of Leopardus (small Felids such as the ocelot and margay)
References and Notes
Further Information
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