Key innovation

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In evolutionary biology, a key innovation, also known as an adaptive breakthrough or key adaptation, is a novel phenotypic trait that allows subsequent radiation and success of a taxonomic group. Typically they bring new abilities that allows the taxa to rapidly diversify and invade niches that were not previously available. The phenomenon helps to explain how some taxa are much more diverse and have many more species than their sister taxa. The term was first used in 1949 by Alden H. Miller who defined it as "key adjustments in the morphological and physiological mechanism which are essential to the origin of new major groups", [1] although a broader, contemporary definition holds that "a key innovation is an evolutionary change in individual traits that is causally linked to an increased diversification rate in the resulting clade". [2]

Contents

The theory of key innovations has come under attack because it is hard to test in a scientific manner, but there is evidence to support the idea. [3]

Mechanism

The mechanism by which a key innovation leads to taxonomic diversity is not certain but several hypotheses have been suggested: [2]

Increasing individual fitness

A key innovation may, by increasing the fitness of individuals of the species, result in extinction becoming less likely and allow the taxa to expand and speciate. [2]

Latex and resin canals in plants are used to deter predators by releasing a sticky secretion when punctured which can immobilise insects and some contain toxic or foul tasting substances. They have evolved independently approximately 40 times and are considered a key innovation. By increasing the plant's resistance to predation the canals increase the species fitness and allow them to escape being eaten, at least until the predator evolves an ability to overcome the defence. During the period of resistance the plants are less likely to become extinct and can diversify and speciate, and as such taxa with latex and resin canals are more diverse than their canal lacking sister taxa. [2]

Novel niche invasion

A human molar with four cusps Lower wisdom tooth.jpg
A human molar with four cusps

A key innovation may allow a species to invade a new region or niche and thus be freed from competition, allowing subsequent speciation and radiation.

A classic example of this is the fourth cusp of mammalian molars, the hypocone, which allowed early mammalian ancestors to effectively digest their generalised diet. The precursors to this, the triconodont teeth of reptiles, were adapted for gripping and slicing rather than chewing. The evolution of the hypocone and flat molars later allowed animals to adapt to a herbivorous diet as they could be used to break down tough plant matter through grinding. The evolution of this ability led to mammals being able to adapt to utilise a huge variety of food sources, [4] and allowed early mammals to invade novel niches through the evolution of specialised herbivores, which experienced relative success during the middle eocene. Specialising for a plant based diet offered early herbivores sufficient resources to radiate as energy was not lost to higher trophic levels and few competitors existed at the time. [4]

Reproductive isolation

The long nectar spurs on this Aquilegia allow specialisation for a certain pollinator. Aquilegia columbine magpie cultivar 2.jpg
The long nectar spurs on this Aquilegia allow specialisation for a certain pollinator.

A key innovation may result in reproductive isolation, whereby those individuals with the innovation no longer breed with those without. This can lead to rapid speciation as the two populations separate and accumulate mutations.

The nectar spurs in Aquilegia , a diverse genus of flowering plant, are considered a key innovation because of this. Nectar spurs aid in pollination by making the nectar further from the stamen, ensuring that insect or bird pollinators pick up pollen as they access it. These led rapid speciation within the genus as plants and their pollinators can become specialised to each other i.e. a species of pollinator exclusively feeds from a species of plant, and thus plant populations could easily become reproductively isolated from one another. In addition the shape and size of the nectar spur can evolve in response to pollinator adaptations, developing a co-evolutionary relationship. The genus Aquilegia has over 50 species. [3]

Criticisms

As an evolutionary theory, key innovations has come under critical scrutiny due to the fact that it is hard to test. Identification depends on finding correlation between the innovation and increased diversity by comparing sister taxa, but this does not prove causality or isolate other causes of diversity such as stochasticity or habitat, and it is possible to 'cherry pick' examples that fit the hypothesis. [5] In addition the retrospective identification of key innovations offers little in terms of understanding the processes and pressures that resulted in the adaptation, and may identify a very complex evolutionary process as a single event. An example of this is the evolution of avian flight, which was identified as a key innovation in 1963 by Ernst Mayr. [6] However, separate evolutionary changes had to occur throughout the physiology of the avian ancestor, including the enlargement of the cerebellum and the enlargement and ossification of the sternum. These adaptations arose separately, and millions of years apart, [5] not in one step.

See also

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Macroevolution Evolution on a scale at or above the level of species

Macroevolution in the modern sense is evolution that is guided by selection among interspecific variation, as opposed to selection among intraspecific variation in microevolution. This modern definition differs from the original concept, which referred macroevolution to the evolution of taxa above the species level.

Coevolution Two or more species influencing each others evolution

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Index of evolutionary biology articles

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Peripatric speciation Speciation in which a new species is formed from an isolated smaller peripheral population

Peripatric speciation is a mode of speciation in which a new species is formed from an isolated peripheral population. Since peripatric speciation resembles allopatric speciation, in that populations are isolated and prevented from exchanging genes, it can often be difficult to distinguish between them. Nevertheless, the primary characteristic of peripatric speciation proposes that one of the populations is much smaller than the other. The terms peripatric and peripatry are often used in biogeography, referring to organisms whose ranges are closely adjacent but do not overlap, being separated where these organisms do not occur—for example on an oceanic island compared to the mainland. Such organisms are usually closely related ; their distribution being the result of peripatric speciation.

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Character displacement

Character displacement is the phenomenon where differences among similar species whose distributions overlap geographically are accentuated in regions where the species co-occur, but are minimized or lost where the species' distributions do not overlap. This pattern results from evolutionary change driven by biological competition among species for a limited resource. The rationale for character displacement stems from the competitive exclusion principle, also called Gause's Law, which contends that to coexist in a stable environment two competing species must differ in their respective ecological niche; without differentiation, one species will eliminate or exclude the other through competition.

Ornithophily Pollination by birds

Ornithophily or bird pollination is the pollination of flowering plants by birds. This sometimes coevolutionary association is derived from insect pollination (entomophily) and is particularly well developed in some parts of the world, especially in the tropics, Southern Africa, and on some island chains. The association involves several distinctive plant adaptations forming a "pollination syndrome". The plants typically have colourful, often red, flowers with long tubular structures holding ample nectar and orientations of the stamen and stigma that ensure contact with the pollinator. Birds involved in ornithophily tend to be specialist nectarivores with brushy tongues and long bills, that are either capable of hovering flight or light enough to perch on the flower structures.

Cusp (anatomy)

A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth. The concept is also used with regard to the valve between the right atrium and the right ventricle in the human heart. This valve is closed during ventricular contraction by the tricuspid valve, so named because it usually consists of three cusps or leaflets.

Plant evolution subset of evolutionary phenomena that concern plants

Plant evolution is the subset of evolutionary phenomena that concern plants. Evolutionary phenomena are characteristics of populations that are described by averages, medians, distributions, and other statistical methods. This distinguishes plant evolution from plant development, a branch of developmental biology which concerns the changes that individuals go through in their lives. The study of plant evolution attempts to explain how the present diversity of plants arose over geologic time. It includes the study of genetic change and the consequent variation that often results in speciation, one of the most important types of radiation into taxonomic groups called clades. A description of radiation is called a phylogeny and is often represented by type of diagram called a phylogenetic tree.

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Ecological fitting

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Escape and radiate coevolution

Escape and radiate coevolution is a hypothesis proposing that a coevolutionary 'arms-race' between primary producers and their consumers contributes to the diversification of species by accelerating speciation rates. The hypothesized process involves the evolution of novel defenses in the host, allowing it to "escape" and then "radiate" into differing species.

Ecological speciation

Ecological speciation is a form of speciation arising from reproductive isolation that occurs due to an ecological factor that reduces or eliminates gene flow between two populations of a species. Ecological factors can include changes in the environmental conditions in which a species experiences, such as behavioral changes involving predation, predator avoidance, pollinator attraction, and foraging; as well as changes in mate choice due to sexual selection or communication systems. Ecologically-driven reproductive isolation under divergent natural selection leads to the formation of new species. This has been documented in many cases in nature and has been a major focus of research on speciation for the past few decades.

Outline of evolution Hierarchical outline list of articles related to evolution

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Nectar spur

A nectar spur is a hollow extension of a part of a flower. The spur may arise from various parts of the flower: the sepals, petals, or hypanthium, and often contain tissues that secrete nectar (nectaries). Nectar spurs are present in many clades across the angiosperms, and are often cited as an example of convergent evolution.

This glossary of evolutionary biology is a list of definitions of terms and concepts used in the study of evolutionary biology, population biology, speciation, and phylogenetics, as well as sub-disciplines and related fields. For additional terms from related glossaries, see Glossary of genetics, Glossary of ecology, and Glossary of biology.

Nonadaptive radiations are a subset of evolutionary radiations that are characterized by diversification that is not driven by resource partitioning. The species that are a part of a nonadaptive radiation will tend to have very similar niches, and in many cases will be morphologically similar. Nonadaptive radiations are driven by nonecological speciation. In many cases, this nonecological speciation is allopatric, and the organisms are dispersal-limited such that populations can be geographically isolated within a landscape with relatively similar ecological conditions. For example, Albinaria land snails on islands in the Mediterranean and Batrachoseps salamanders from California each include relatively dispersal-limited, and closely related, ecologically similar species often have minimal range overlap, a pattern consistent with allopatric, nonecological speciation. In other cases, such as certain damselflies and crickets from Hawaii, there can be range overlap in closely related species, and it is likely that sexual selection plays a role in maintaining species boundaries.

References

  1. Miller, Alden (November 22, 1949). "Some ecologic and morphologic considerations in the evolution of higher taxonomic categories". Ornithologie als Biologische Wissenschaft. pp. 84–88.
  2. 1 2 3 4 Heard, S.B.; Hauser, D.L. (1995). "Key evolutionary innovations and their ecological mechanisms". Historical Biology. 10 (2): 151–173. doi:10.1080/10292389509380518.
  3. 1 2 Hodges, S.A. Arnold, M.L. (1995). "Spurring plant diversification: are floral nectar spurs a key innovation?". Proceedings: Biological Sciences. 262 (1365): 343–348. doi:10.1098/rspb.1995.0215. S2CID   86823646.CS1 maint: multiple names: authors list (link)
  4. 1 2 Hunter, J.P. Jernvall, J. (1995). "The hypocone as a key innovation in mammalian evolution". Proceedings of the National Academy of Sciences. 92 (23): 10718–10722. Bibcode:1995PNAS...9210718H. doi:10.1073/pnas.92.23.10718. PMC   40683 . PMID   7479871.CS1 maint: multiple names: authors list (link)
  5. 1 2 Cracraft, Joel (1990). "The origin of evolutionary novel pattern and process at different hierarchical levels". Evolutionary innovations. pp. 21–46.
  6. Mayr, Ernst (1963). Animal Species and Evolution . Cambridge: Harvard University Press. ISBN   0-674-03750-2.