Modular Evolution
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Mosaic evolution (or modular evolution) is the concept, mainly from
palaeontology Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geo ...
, that
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
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 comprises the evolutionary processes and patterns which occur at and above the species level. In contrast, microevolution is evolution occurring within the population(s) of a single species. In other words, microevolution is the ...
.


Background

In the neodarwinist
theory of evolution Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certai ...
, as postulated by
Stephen Jay Gould Stephen Jay Gould ( ; September 10, 1941 – May 20, 2002) was an American Paleontology, paleontologist, Evolutionary biology, evolutionary biologist, and History of science, historian of science. He was one of the most influential and widely re ...
, 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 In evolutionary developmental biology, heterochrony is any genetically controlled difference in the timing, rate, or duration of a developmental process in an organism compared to its ancestors or other organisms. This leads to changes in the s ...
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 in modern humans is more signi ...
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 ovum, egg to adult. The term can also be used to refer to t ...
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 in modern humans is more signif ...
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 ''Homo sapiens'' is a distinct species of the hominid family of primates, which also includes all the great apes. Over their evolutionary history, humans gradually developed traits such as bipedalism, dexterity, and complex language, as well as ...
: the early evolution of
bipedalism Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
in
Australopithecine The australopithecines (), formally Australopithecina or Hominina, are generally any species in the related genera of ''Australopithecus'' and ''Paranthropus''. It may also include members of '' Kenyanthropus'', ''Ardipithecus'', and '' Praeant ...
s, and its modification of the
pelvic girdle The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the Ilium (bone) ...
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 (''archaîos''), 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 who specialized in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stor ...
compared ''Archaeopteryx'' with a small
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
, ''
Compsognathus ''Compsognathus'' (; Ancient Greek, Greek ''kompsos''/κομψός; "elegant", "refined" or "dainty", and ''gnathos''/γνάθος; "jaw") is a genus of small, bipedalism, bipedal, carnivore, carnivorous theropoda, theropod dinosaur. Members o ...
''. Both fossils came from the
Solnhofen Limestone The Solnhofen Limestone or Solnhofen Plattenkalk, formally known as the Altmühltal Formation, is a Jurassic Konservat-Lagerstätte that preserves a rare assemblage of fossilized organisms, including highly detailed imprints of soft bodied organi ...
in
Bavaria Bavaria, officially the Free State of Bavaria, is a States of Germany, state in the southeast of Germany. With an area of , it is the list of German states by area, largest German state by land area, comprising approximately 1/5 of the total l ...
. 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 an exa ...
s of ''Archaeopteryx''. His interest was in the basic affinity of birds and reptiles, which he united as the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
''
Sauropsida Sauropsida (Greek language, Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the Class (biology), class Reptile, Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern repti ...
''. The peculiarity here is that the rest of the skeleton had not changed. * Meadow voles during the last 500,000 years. * The
pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
''
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 Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
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 Rhamphorhync ...
) and short-tailed (
pterodactyloid Pterodactyloidea ( ; derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger") is one of the two traditional suborders of pterosaurs ("wing lizards"), and contai ...
) 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 is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
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 being a classic example of adaptive radiation and for their remarkable diversity in beak form and function. They ...
, it can also be seen in the evolutionary process of
hominin The Hominini (hominins) form a taxonomic tribe of the subfamily Homininae (hominines). They comprise two extant genera: ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos), and in standard usage exclude the genus '' Gorilla'' ( gorillas) ...
. 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 Fossil Site, Cradle of Humankind, Malapa Cave, Cradle of Humankind, South Africa. It is known from a partial juvenile skeleton, the holotype MH1, and a ...
'', ''
H. naledi '' Homo naledi'' is an extinct species of archaic human discovered in 2013 in the Rising Star Cave system, Gauteng province, South Africa, part of the Cradle of Humankind, dating back to the Middle Pleistocene 335,000–236,000 years ago. The ...
'', and ''
H. floresiensis ''Homo floresiensis'' , also known as "Flores Man" or "Hobbit" (after the fictional species), is an extinct species of small archaic humans that inhabited the island of Flores, Indonesia, until the arrival of modern humans about 50,000 years ...
''. 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 that consists of grammar and vocabulary. It is the primary means by which humans convey meaning, both in spoken and signed language, signed forms, and may also be conveyed through writing syste ...
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 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. Measuring brain size and cranial capacity is relevant both to humans and other animals, and can be done by wei ...
. This is provided and shown by
Robin Dunbar Robin Ian MacDonald Dunbar (born 28 June 1947) is a British biological anthropologist, evolutionary psychologist, and specialist in primate behaviour. Dunbar is professor emeritus of evolutionary psychology of the Social and Evolutionary Neuros ...
'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'' (, ; or (, ) is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera ''Homo'' (which includes modern humans), ''Paranthropus'', and ''Kenyanthropus'' evolved from some ''Aus ...
'' 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 a ...
*
Coextinction Coextinction and cothreatened refer to the phenomenon of the loss or decline of a host species resulting in the loss or endangerment of another species that depends on it, potentially leading to cascading effects across trophic levels. The term w ...
*
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 comm ...
*
Evolutionary developmental biology Evolutionary developmental biology, informally known as evo-devo, is a field of biological research that compares the developmental biology, developmental processes of different organisms to infer how developmental processes evolution, evolved. ...
*
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 pa ...


References

{{Evolution Evolutionary biology concepts