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.[1][2]

Parallel vs. convergent evolution

Evolution at an amino acid position. In each case, the left-hand species changes from incorporating alanine (A) at a specific position within a protein in a hypothetical common ancestor deduced from comparison of sequences of several species, and now incorporates serine (S) in its present-day form. The right-hand species may undergo divergent evolution (alanine replaced with threonine instead), parallel evolution (alanine also replaced with serine), or convergent evolution (threonine replaced with serine) at this amino acid position relative to that of the first species.

Given a particular trait that occurs in each of two lineages descended from a specified ancestor, it is possible in theory to define parallel and convergent evolutionary trends strictly, and distinguish them clearly from one another.[2] However the criteria for defining convergent as opposed to parallel evolution often are unclear in practice, so that arbitrary diagnosis is common in some cases.

When two species are similar in a particular character, evolution is defined as parallel if the ancestors shared that similarity; if they did not, the evolution of that character in those species is defined as convergent. However, this distinction is not clear-cut. For one thing, the stated conditions are partly a matter of degree; all organisms share more or less recent common ancestors. In evolutionary biology the question of how far back to look for similar ancestors, and how similar those ancestors need to be for one to consider parallel evolution to have taken place, cannot always be resolved. Some scientists accordingly have argued that parallel evolution and convergent evolution are more or less indistinguishable.[3] Others insist that in practice we should not shy away from the gray area because many important distinctions between parallel and convergent evolution remain.[4]

When the ancestral forms are unspecified or unknown, or the range of traits considered is not clearly specified, the distinction between parallel and convergent evolution becomes more subjective. For instance, Richard Dawkins in The Blind Watchmaker describes the striking similarity between placental and marsupial forms as the outcome of convergent evolution, because mammals on their respective ancestral continents had a long evolutionary history before the extinction of the dinosaurs. That period of separation would have permitted the accumulation of many relevant differences. Stephen Jay Gould differed; he described some of the same examples as having started from the common ancestor of all marsupials and placentals, and hence amounting to parallel evolution. And certainly, whenever similarities can be described in concept as having evolved from a common attribute deriving from a single remote ancestral line, that legitimately may be regarded as parallel evolution.

In contrast, where quite different structures clearly have been co-opted to a similar form and function, one must necessarily regard the evolution as convergent. For example, consider Mixotricha paradoxa, a eukaryotic microbe which has assembled a system of rows of apparent cilia and basal bodies closely resembling the system in ciliates. However, on inspection it turns out that in Mixotricha paradoxa, what appear to be cilia actually are smaller symbiont microorganisms; there is no question of parallel evolution in such a case. Again, the differently oriented tails of fish and whales derived at vastly different times from radically different ancestors and any similarity in the resultant descendants must therefore have evolved convergently; any case in which lineages do not evolve together at the same time in the same ecospace might be described as convergent evolution at some point in time.

The definition of a trait is crucial in deciding whether a change is seen as divergent, or as parallel or convergent. For example, the evolution of the sesamoid "thumb" of the giant panda certainly is not parallel to that of the thumbs of primates, particularly hominins, and it also differs morphologically from primate thumbs, but from some points of view it might be regarded as convergent in function and appearance.

Again, in the image above, note that since serine and threonine possess similar structures with an alcohol side chain, the example marked "divergent" would be termed "parallel" if the amino acids were grouped by similarity instead of being considered individually. As another example, if genes in two species independently become restricted to the same region of the animals through regulation by a certain transcription factor, this may be described as a case of parallel evolution - but examination of the actual DNA sequence will probably show only divergent changes in individual basepair positions, since a new transcription factor binding site can be added in a wide range of places within the gene with similar effect.

A similar situation occurs considering the homology of morphological structures. For example, many insects possess two pairs of flying wings. In beetles, the first pair of wings is hardened into elytra, wing covers with little role in flight, while in flies the second pair of wings is condensed into small halteres used for balance. If the two pairs of wings are considered as interchangeable, homologous structures, this may be described as a parallel reduction in the number of wings, but otherwise the two changes are each divergent changes in one pair of wings.

Similar to convergent evolution, evolutionary relay describes how independent species acquire similar characteristics through their evolution in similar ecosystems, but not at the same time, such as the dorsal fins of sharks, cetaceans and ichthyosaurs.

Examples

  • Colouration that serves as a warning to predators and for mating displays has evolved in many different species.
  • In the plant kingdom, the most familiar examples of parallel evolution are the forms of leaves, where very similar patterns have appeared again and again in separate genera and families.
  • In Arabidopsis thaliana it has been suggested that populations adapt to local climate through parallel evolution [5]
  • In butterflies, many close similarities are found in the patterns of wing colouration, both within and among families.
  • Old and New World porcupines shared a common ancestor, both evolved strikingly similar quill structures; this is also an example of convergent evolution as similar structures evolved in hedgehogs, echidnas and tenrecs.
  • Some extinct archosaurs evolved an upright posture and likely were warm-blooded. These two characteristics are also found in most mammals. Modern crocodiles have a four chambered heart and a crurotarsal, the latter being also a characteristic of therian mammals.
  • The extinct pterosaurs and the birds both evolved wings as well as a distinct beak, but not from a recent common ancestor.
  • Internal fertilization has evolved independently in sharks, some amphibians and amniotes.
  • The patagium is a fleshy membrane that is found in gliding mammals such as flying lemurs, flying squirrels, sugar gliders and the extinct Volaticotherium. These mammals all acquired the patagium independently.
  • Pyrotherians evolved a body plan similar to proboscideans.
  • The extinct South American litoptern ungulate Thoatherium had legs that are difficult to distinguish from those of horses.
  • The eye of the octopus has the same complicated structure as the human eye. As a result, it is often substituted in studies of the eye when using a human eye would be inappropriate. As the two species diverged at the time animals evolved into deuterostomes and protostomes this is extraordinary.
  • Certain arboreal frog species, 'flying' frogs, in both Old World families and New World families have developed the ability of gliding flight. They have "enlarged hands and feet, full webbing between all fingers and toes, lateral skin flaps on the arms and legs, and reduced weight per snout-vent length".[6]
  • The tree plant habit has evolved separately in unrelated classes of plants.

Parallel evolution between marsupials and placentals

A number of examples of parallel evolution are provided by the two main branches of the mammals, the placentals and marsupials, which have followed independent evolutionary pathways following the break-up of land-masses such as Gondwanaland roughly 100 million years ago. In South America, marsupials and placentals shared the ecosystem (before the Great American Interchange); in Australia, marsupials prevailed; and in the Old World and North America the placentals won out. However, in all these localities mammals were small and filled only limited places in the ecosystem until the mass extinction of dinosaurs sixty-five million years ago. At this time, mammals on all three landmasses began to take on a much wider variety of forms and roles. While some forms were unique to each environment, surprisingly similar animals have often emerged in two or three of the separated continents. Examples of these include the placental sabre-toothed cats (Machairodontinae) and the South American marsupial sabre-tooth (Thylacosmilus); the Tasmanian wolf and the European wolf; likewise marsupial and placental moles, flying squirrels, and (arguably) mice.

Parallel coevolution of traits between hummingbirds and sunbirds contributing to ecological guilds

Hummingbirds and sunbirds, two nectarivorous bird lineages in the New and Old Worlds have parallelly evolved a suite of specialized behavioral and anatomical traits. These traits (bill shape, digestive enzymes, and flight) allow the birds to optimally fit the flower-feeding-and-pollination ecological niche they occupy, which is shaped by the birds' suites of parallel traits. Thus, a parallel coevolved behavioral syndrome within the birds creates an emergent guild of highly specialized birds and highly adapted plants, each exploiting the other's involvement in the flowers' pollination in the Old World and New World alike.[7]

Bill morphology

The bill shape of nectarivores, being long and needle-like, allows them to reach down a flower’s pistil/stamen and get at the nectar within. Nectarivores may also use their specialized bills to engage in nectar robbing, a practice seen in both hummingbirds and sunbirds in which the bird gets nectar by making a hole in the base of the flower's corolla tube instead of inserting its bill through the tube as is standard, thus "robbing" the flower of nectar since it is not pollinated it in return.[8]

Digestive morphology

The capacity of nectarivores to digest sucrose is far greater than that of other avian taxa. This difference is due to an analogous high concentration of sucrase-isomaltase, an enzyme that hydrolyzes sucrose. Sucrase activity per unit intestinal surface area appears to be higher in nectarivores than in other birds, meaning these nectarivorous avians can digest more sucrose than other taxa and in a quicker manner because the enzymes hydrolyzing sucrose are more abundant in their digestive tract.[9] Moreover, the Adaptive Modulation Hypothesis[10] does not apply for nectarivores and sugar-digesting enzymes, meaning that two lineages of nectarivores should not necessarily both have high sucrase-isomaltase concentrations even though they both eat nectar. Thus, parallel acquisition of analogous sucrose digestive capability is a reasonable conclusion because there is no apparent cause for the two lineages sharing this high enzyme concentration, even when the similarities of their diets are considered. In other words, specialized nectarivory is not necessarily associated with a high capacity to digest sucrose, meaning that the specialized sunbirds' and hummingbirds' sucrose digestive capacity is parallel because it is not required, yet both lineages possess it.

Mutualism and guilds

Nectarivores and ornithophilous flowers often exist in mutualistic guild relationships facilitated by the bird's bill shape, food source, and digestive ability acting in concert with the flower's tube shape and adaptation to pollination by hovering or perching birds.[11] The birds eat nectar using their long, thin bills and, in so doing, collect pollen on their bills; this pollen is then transferred to the next flower they feed on. This mutualism coevolved in parallel between the Old World and New World birds and their respective flowers.

Moreover, the digestive enzyme activity in nectarivores matching the nectar composition in their respective flowers appears to be an example of parallel coevolution between plants and pollinators across continents, as the nectarivorous lineages independently evolved the ability to digest the nectar specific to their flowers, resulting in distinct guilds that evolved via the same means.[11][9]

References

  1. Parallel evolution, an example may be the Pyrotherians evolved a body plan similar to proboscideans: Online Biology Glossary Archived 2007-07-13 at the Wayback Machine
  2. Zhang, J. and Kumar, S. 1997. Detection of convergent and parallel evolution at the amino acid sequence level Archived 2016-03-03 at the Wayback Machine. Mol. Biol. Evol. 14, 527-36.
  3. ARENDT, J; REZNICK, D (January 2008). "Convergence and parallelism reconsidered: what have we learned about the genetics of adaptation?". Trends in Ecology & Evolution. 23 (1): 26–32. doi:10.1016/j.tree.2007.09.011. PMID 18022278.
  4. Pearce, T. (10 November 2011). "Convergence and Parallelism in Evolution: A Neo-Gouldian Account". The British Journal for the Philosophy of Science. 63 (2): 429–448. doi:10.1093/bjps/axr046.
  5. Stearns, F. W. and C. B. Fenster (2013). Evidence for parallel adaptation across the natural range of Arabidopsis thaliana. Ecology & Evolution. http://onlinelibrary.wiley.com/doi/10.1002/ece3.622/full
  6. Emerson, S.B., & Koehl, M.A.R. (1990). "The interaction of behavioral and morphological change in the evolution of a novel locomotor type: 'Flying' frogs." Evolution, 44(8), 1931-1946
  7. Janeček, Štěpán; Chmel, Kryštof; Uceda Gómez, Guillermo; Janečková, Petra; Chmelová, Eliška; Sejfová, Zuzana; Luma Ewome, Francis (February 2020). "Ecological fitting is a sufficient driver of tight interactions between sunbirds and ornithophilous plants". Ecology and Evolution. 10 (4): 1784–1793. doi:10.1002/ece3.5942. ISSN 2045-7758. PMC 7042734. PMID 32128116.
  8. Juan Francisco Ornelas. Serrate Tomia: An Adaptation for Nectar Robbing in Hummingbirds?. The Auk, Volume 111, Issue 3, Januar 1994, Pages 703710.
  9. McWhorter, Todd J.; Rader, Jonathan A.; Schondube, Jorge E.; Nicolson, Susan W.; Pinshow, Berry; Fleming, Patricia A.; Gutiérrez-Guerrero, Yocelyn T.; Martínez del Rio, Carlos (July 2021). "Sucrose digestion capacity in birds shows convergent coevolution with nectar composition across continents". iScience. 24 (7): 102717. doi:10.1016/j.isci.2021.102717. ISSN 2589-0042. PMC 8246590. PMID 34235412.
  10. Karasov, W. H. (1992-09-01). "Tests of the adaptive modulation hypothesis for dietary control of intestinal nutrient transport". American Journal of Physiology-Regulatory, Integrative and Comparative Physiology. 263 (3): R496–R502. doi:10.1152/ajpregu.1992.263.3.R496. ISSN 0363-6119.
  11. Janeček, Štěpán; Bartoš, Michael; Njabo, Kevin Yana (2015-01-22). "Convergent evolution of sunbird pollination systems of Impatiens species in tropical Africa and hummingbird systems of the New World". Biological Journal of the Linnean Society. 115 (1): 127–133. doi:10.1111/bij.12475. ISSN 0024-4066.
Notes
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