Evolutionary anachronism
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Evolutionary anachronism is a concept in
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
named by Connie C. Barlow in her book, ''The Ghosts of Evolution'' (2000). It refers to attributes of living species that are best explained as a result of having been favorably selected in the past due to
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
with other biological species that have since become
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. When this context is removed, the natural attributes appear as unexplained energy investments by the living organism, with no apparent benefit, and perhaps are prejudicial to the continued reproduction of the surviving species. The general theory was formulated by
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
n-based American
botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
Daniel Janzen Daniel Hunt Janzen (born January 18, 1939 in Milwaukee, Wisconsin) is an American evolutionary ecologist, and conservationist. He divides his time between his professorship in biology at the University of Pennsylvania, where he is the DiMaura ...
and
University of Arizona The University of Arizona (Arizona, U of A, UArizona, or UA) is a Public university, public Land-grant university, land-grant research university in Tucson, Arizona. Founded in 1885 by the 13th Arizona Territorial Legislature, it was the first ...
-based
geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, althou ...
Paul S. Martin (a prominent defender of the overkill hypothesis to explain the
Quaternary extinction event The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...
) in a ''
Science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence ...
'' article published in 1982, titled ''Neotropical Anachronisms: The fruit the
gomphothere Gomphotheres are any members of the diverse, extinct taxonomic family Gomphotheriidae. Gomphotheres were elephant-like proboscideans, but do not belong to the family Elephantidae. They were widespread across Afro-Eurasia and North America dur ...
s ate.'' Previously, in 1977,
Stanley Temple Stanley A. "Stan" Temple is an American avian ecologist and wildlife biologist. He is the Beers-Bascom Professor in Conservation, Professor of Wildlife Ecology and Professor of Environmental Studies at the University of Wisconsin–Madison. At Cor ...
had proposed a similar idea to explain the decline of the
Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label= Mauritian Creole, Moris ), officially the Republic of Mauritius, is an island nation in the Indian Ocean about off the southeast coast of the African continent, east of Madagascar. It ...
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
tree
tambalacoque ''Sideroxylon grandiflorum'', known as tambalacoque or dodo tree, is a long-lived mesocaul tree in the sapote family Sapotaceae, superfiially resembling the unrelated Plumeria, but the Dodo Tree's flowers and fruit are cauliflorous. endemic t ...
following the extinction of the iconic
dodo The dodo (''Raphus cucullatus'') is an extinct flightless bird that was endemic to the island of Mauritius, which is east of Madagascar in the Indian Ocean. The dodo's closest genetic relative was the also-extinct Rodrigues solitaire. ...
. Janzen, Martin and Barlow mainly discussed evolutionary anachronisms in the context of
seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vector ...
and passive defense strategies exhibited by plants that had evolved alongside disappeared megaherbivores, although some examples apply to animal species as relict behavior. Evolutionary anachronisms are not the same as
vestigiality Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
. Though both concepts refer ultimately to organs that evolved in response to environmental pressures not currently present, in the case of anachronisms, the original function of the organ and the capacity of the organism to use it are retained. For example, the present-day absence of
avocado The avocado (''Persea americana'') is a medium-sized, evergreen tree in the laurel family ( Lauraceae). It is native to the Americas and was first domesticated by Mesoamerican tribes more than 5,000 years ago. Then as now it was prized for ...
-eating
gomphothere Gomphotheres are any members of the diverse, extinct taxonomic family Gomphotheriidae. Gomphotheres were elephant-like proboscideans, but do not belong to the family Elephantidae. They were widespread across Afro-Eurasia and North America dur ...
s does not render avocado pulp vestigial, rudimentary or incapable of its original function in seed nutrition and dispersal ( zoochory) if a new suitable ecological partner were to appear.


Megafauna dispersal syndrome

Dispersal syndromes are complexes of fruit traits that enable plants to disperse seeds. The kind of fruits that birds are attracted to are usually small, with only a thin protective skin, and the colors are red or dark shades of blue or purple. Fruits categorized as mammal syndrome are bigger than bird fruits. They possess a tough rind or husk, emit a strong odor when ripe but retain a dull coloration of brown, burnished yellow, orange or remain green, because most mammals have a powerful sense of smell but poor color vision in general,
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter including ...
being the most notable exception. The
megafauna In terrestrial zoology, the megafauna (from Greek μέγας ''megas'' "large" and New Latin ''fauna'' "animal life") comprises the large or giant animals of an area, habitat, or geological period, extinct and/or extant. The most common thresho ...
dispersal syndrome refers to those attributes of fruits that evolved in order to attract megafauna (animals that weigh or weighed more than 44 kilograms) as primary dispersal agents. Since the
Holocene extinction The Holocene extinction, or Anthropocene extinction, is the ongoing extinction event during the Holocene epoch. The extinctions span numerous families of bacteria, fungi, plants, and animals, including mammals, birds, reptiles, amphibians, fish, ...
, large herbivores have become extinct outside
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and to a lesser extent
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, leaving these fruits without a suitable dispersal mechanism in the absence of
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people ...
.


Common megafaunal dispersal traits

*Large fruit, best suited to be consumed whole by large animals without seed loss. *Fruit grows on or close to the trunk, or on stout branches. * Indehiscent fruit that retains its seeds upon ripening. *Seeds deter or elude being ground up by teeth through having a thick, tough or hard
endocarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Aggr ...
; or bitter, peppering or nauseating toxins. They are also difficult to separate from the pulp, which is tasty and soft, to deter seed spitting. *The seeds benefit from—or even require—physical or chemical abrasion to germinate. *If tropical, the fruit drops on or just before ripening, stopping
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
s from eating them. In colder climates, the fruit stays on the branch for a prolonged time, keeping it away from predation by ineffectual seed dispersers like
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s. *"Looks, feels, smells, and tastes" like other fruits known to be dispersed by megafauna where megafauna still exists.


Ecological indicators of missing dispersal partners

*The fruit either rots where it falls or is ineffectually disseminated by current dispersal agents. *The plant is more common where
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to ani ...
(proxy for megafauna) are present. *The seeds germinate and grow well in upland habitats where planted, but the species almost exclusively inhabits floodplains (where water flow disperses the seeds) in the wild. *The geographic range is inexplicably patchy or restricted.


Proposed examples in plants


Afrotropical realm


Madagascar


Australasian realm


New Zealand


Indomalayan realm


Nearctic realm


Neotropical realm


Oceanian realm


Palearctic realm


Proposed examples in animals


In popular culture

The phenomenon was referenced (though not by name) in the 1259 issue "Bee Orchid" of the online comic strip ''
xkcd ''xkcd'', sometimes styled ''XKCD'', is a webcomic created in 2005 by American author Randall Munroe. The comic's tagline describes it as "a webcomic of romance, sarcasm, math, and language". Munroe states on the comic's website that the name ...
'' by
Randall Munroe Randall Patrick Munroe (born October 17, 1984) is an American cartoonist, author, and engineer best known as the creator of the webcomic ''xkcd''. Munroe has worked full-time on the comic since late 2006. In addition to publishing a book of th ...
, published on September 2, 2013. In the comic, it is claimed that the orchid '' Ophrys apifera'' mimics the female of a bee species to attract males and ensure pollination, but that the bee is extinct and the orchid is doomed to follow, only delaying it by resorting to less effective
self-pollination Self-pollination is a form of pollination in which pollen from the same plant arrives at the stigma of a flower (in flowering plants) or at the ovule (in gymnosperms). There are two types of self-pollination: in autogamy, pollen is transferr ...
. In reality, only the first part is true: the flower mimics a bee, and reproduces exclusively by self-pollination in northern Europe, but it is pollinated successfully by bees of the genus ''
Eucera ''Eucera'' is a genus of bees in the family Apidae, subfamily Apinae, and tribe Eucerini – the long-horned bees. Description As in most members of the tribe Eucerini, the antennae of males are very long. Old World ''Eucera'' can be identifi ...
'' in the Mediterranean region.


See also

*
Coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
*
Coextinction Coextinction and cothreatened refer to the phenomena of the loss or decline of a host species resulting in the loss or endangerment of an other species that depends on it, potentially leading to cascading effects across trophic levels. The term or ...
* Evolutionary arms race *
Evolutionary mismatch Evolutionary mismatch, also known as mismatch theory or evolutionary trap, is a concept in evolutionary biology that refers to evolved traits that were once advantageous but became maladaptive due to changes in the environment. This can take pla ...
* Evolutionary trap *
Holocene extinction The Holocene extinction, or Anthropocene extinction, is the ongoing extinction event during the Holocene epoch. The extinctions span numerous families of bacteria, fungi, plants, and animals, including mammals, birds, reptiles, amphibians, fish, ...
*
Keystone species A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintaini ...
*
Pleistocene extinction The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...
* Pleistocene rewilding


References

{{Reflist, 30em


External links


Youtube video "Why are Lemurs Terrified of Predators that don't Exist?"Gourds and squashes (Cucurbita spp.) adapted to megafaunal extinction and ecological anachronism through domestication
Evolutionary biology Ecology Habitat