Mosaic Evolution
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Mosaic evolution (or modular evolution) is the concept, mainly from
palaeontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
, that
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary change takes place in some body parts or systems without simultaneous changes in other parts. Another definition is the "evolution of characters at various rates both within and between species".Carroll R.L. 1997. ''Patterns and processes of vertebrate evolution''. Cambridge University Press. 408 Its place in evolutionary theory comes under long-term trends or
macroevolution Macroevolution usually means the evolution of large-scale structures and traits that go significantly beyond the intraspecific variation found in microevolution (including speciation). In other words, macroevolution is the evolution of taxa abov ...
.


Background

In the neodarwinist
theory of evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
, as postulated by
Stephen Jay Gould Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Gould sp ...
, there is room for differing development, when a life form matures earlier or later, in shape and size. This is due to allomorphism. Organs develop at differing rhythms, as a creature grows and matures. Thus a " heterochronic clock" has three variants: 1) time, as a straight line; 2) general size, as a curved line; 3) shape, as another curved line. When a creature is advanced in size, it may develop at a smaller rate. Alternatively, it may maintain its original size or, if delayed, it may result in a larger sized creature. That is insufficient to understand heterochronic mechanism. Size must be combined with shape, so a creature may retain
paedomorphic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
features if advanced in shape or present recapitulatory appearance when retarded in shape. These names are not very indicative, as past theories of development were very confusing. Gould, S.J. 1977. ''Ontogeny and phylogeny''. Belknap Press of Harvard University Press. A creature in its
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
may combine heterochronic features in six vectors, although Gould considers that there is some binding with growth and sexual maturation. A creature may, for example, present some
neotenic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
features and retarded development, resulting in new features derived from an original creature only by regulatory genes. Most novel human features (compared to closely related apes) were of this nature, not implying major change in structural genes, as was classically considered.


Taxonomic range

It is not claimed that this pattern is universal, but there is now a wide range of examples from many different taxa, including: *
Hominid evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of ''Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual development of ...
: the early evolution of
bipedalism Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
in
Australopithecine Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally ''Australopithecus'' (cladistically including the genus, genera ''Homo'', ''Paranthropus'', and ''Kenyanthropus''), and it typically in ...
s, and its modification of the
pelvic girdle The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The p ...
took place well before there was any significant change in the skull, or brain size.Stanley, S.M. 1979. ''Macroevolution: pattern and process''. Freeman, San Francisco. p154 Jurmain, Robert. et al. 2008. ''Introduction to Physical Anthropology''. Thompson Wadsworth. p479 *''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
''. Nearly 150 years ago,
Thomas Henry Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The storie ...
compared ''Archaeopteryx'' with a small
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
, ''
Compsognathus ''Compsognathus'' (; Greek ''kompsos''/κομψός; "elegant", "refined" or "dainty", and ''gnathos''/γνάθος; "jaw") is a genus of small, bipedal, carnivorous theropod dinosaur. Members of its single species ''Compsognathus longipes'' c ...
''. Both fossils came from the Solnhofen Limestone in
Bavaria Bavaria ( ; ), officially the Free State of Bavaria (german: Freistaat Bayern, link=no ), is a state in the south-east of Germany. With an area of , Bavaria is the largest German state by land area, comprising roughly a fifth of the total lan ...
. Huxley showed that the two were very similar, except for the front limbs and
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s of ''Archaeopteryx''. His interest was in the basic affinity of birds and reptiles, which he united as the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
''
Sauropsida Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
''. The peculiarity here is that the rest of the skeleton had not changed. * Meadow voles during the last 500,000 years. * The
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
''
Darwinopterus ''Darwinopterus'' (meaning "Darwin's wing") is a genus of pterosaur, discovered in China and named after biologist Charles Darwin. Between 30 and 40 fossil specimens have been identified, all collected from the Tiaojishan Formation, which dates t ...
''. The type species, ''D. modularis'' was the first known pterosaur to display features of both long-tailed (
rhamphorhynchoid The Rhamphorhynchoidea forms one of the two suborders of pterosaurs and represents an evolutionary grade of primitive members of flying reptiles. This suborder is paraphyletic unlike the Pterodactyloidea, which arose from within the Rhamphorhynch ...
) and short-tailed (
pterodactyloid Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...
) pterosaurs. *
Evolution of the horse The evolution of the horse, a mammal of the family Equidae, occurred over a geologic time scale of 50 million years, transforming the small, dog-sized, forest-dwelling '' Eohippus'' into the modern horse. Paleozoologists have been able to piece ...
, in which the major changes took place at different times, not all simultaneously.Maynard Smith, John 1993. ''The theory of evolution''. Cambridge University Press. 3rd ed + new Introduction. pp285–290 *
Mammalian evolution The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian (geology), Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synaps ...
, especially during the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
provides a clear and well-understood example.


Mosaic evolution (in hominin)

Although mosaic evolution is usually seen in terms of animals such as
Darwin's finches Darwin's finches (also known as the Galápagos finches) are a group of about 18 species of passerine birds. They are well known for their remarkable diversity in beak form and function. They are often classified as the subfamily Geospizinae or t ...
, it can also be seen in the evolutionary process of
hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
. To help further explain the meaning of mosaic evolution in hominin, mosaicism will get broken down into three subgroups. Group 1 includes related species developing independently, of which carry deep variability in their own morphological structure. Examples of this can be seen within comparisons of ''
A. sediba ''Australopithecus sediba'' is an extinct species of australopithecine recovered from Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a partial adult female skeleton, the par ...
'', ''
H. naledi '' Homo naledi'' is an extinct species of archaic human discovered in 2013 in the Rising Star Cave, Cradle of Humankind, South Africa dating to the Middle Pleistocene 335,000–236,000 years ago. The initial discovery comprises 1,550 specimens ...
'', and ''
H. floresiensis ''Homo floresiensis'' also known as "Flores Man"; nicknamed "Hobbit") is an extinct species of small archaic human that inhabited the island of Flores, Indonesia, until the arrival of modern humans about 50,000 years ago. The remains of an ...
''. Group 2 relies on the different environmental impacts on the changes of a species. An example of this is the variability of bipedalism forming independently within all related species of hominin. Lastly, Group 3 involves the presence of behavior such as the human vernacular.
Language Language is a structured system of communication. The structure of a language is its grammar and the free components are its vocabulary. Languages are the primary means by which humans communicate, and may be conveyed through a variety of met ...
is a mosaic composite of various elements working together for one specific attribute, and this is not a single trait an offspring can inherit directly. In addition, it has been shown that an increase in social interactions corresponds to the
evolution of human intelligence The evolution of human intelligence is closely tied to the evolution of the human brain and to the origin of language. The timeline of human evolution spans approximately seven million years, from the separation of the genus '' Pan'' until the em ...
or in other words, an increase in
brain size The size of the brain is a frequent topic of study within the fields of anatomy, biological anthropology, animal science and evolution. Brain size is sometimes measured by weight and sometimes by volume (via MRI scans or by skull volume). Neur ...
. This is provided and shown by
Robin Dunbar Robin Ian MacDonald Dunbar (born 28 June 1947) is a British anthropologist and evolutionary psychologist and a specialist in primate behaviour. He is currently head of the Social and Evolutionary Neuroscience Research Group in the Department o ...
's social brain hypothesis. Moreover, this can be used as a level of transition in human evolution; of which also includes dental shapes. Brain size has shown intra-specific mosaic variability within its own development, as these differences are a result of environmental limitations. In other words, independent variability of brain structure is seen more when brain regions are unassociated from one another, ultimately, giving rise to perceptible features. When comparing current brain size and capacity between humans and chimpanzees, the ability to predict the evolutionary change between their ancestors was incredibly insightful. This granted the discovery that "local spatial interactions" were the main effect of the limitations. Furthermore, alongside the cranial capacity and structure of the brain, dental shape provides another example of mosaicism. Using fossil record, dental shape showed mosaic evolution within the canine teeth found in early hominin. Reduction of canine sizes are seen as an authentication mark of human ancestor evolution. However, '' A. anamensis'', discovered in Kenya, was found to have the largest mandibular canine root as part of ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australopi ...
'' evolution. This alters the authentication mark because the dimorphism between root and crown reduction has not been assessed. Although canine reduction has probably occurred prior to the evolution of ''Australopithecus'', "changes in canine shape, in both crowns and roots, occurred in a mosaic fashion throughout the ''A. anamensis–afarensis'' lineage".


See also

*
Co-adaptation In biology, co-adaptation is the process by which two or more species, genes or phenotypic traits undergo adaptation as a pair or group. This occurs when two or more interacting characteristics undergo natural selection together in response to the ...
*
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 ...
*
Convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
*
Evolutionary developmental biology Evolutionary developmental biology (informally, evo-devo) is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved. The field grew from 19th-century beginni ...
*
Parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...


References

{{Evolution Evolutionary biology concepts