Convergent evolution

Two succulent plant genera, Euphorbia and Astrophytum, are only distantly related, but the species within each have converged on a similar body form.

E obesa symmetrica ies
Astrophytum asterias1

Convergent evolution is the independent evolution of similar features in species of different lineages. Convergent evolution creates analogous structures that have similar form or function but were not present in the last common ancestor of those groups. The cladistic term for the same phenomenon is homoplasy. The recurrent evolution of flight is a classic example, as flying insects, birds, pterosaurs, and bats have independently evolved the useful capacity of flight. Functionally similar features that have arisen through convergent evolution are analogous, whereas homologous structures or traits have a common origin but can have dissimilar functions. Bird, bat, and pterosaur wings are analogous structures, but their forelimbs are homologous, sharing an ancestral state despite serving different functions.

The opposite of convergence is divergent evolution, where related species evolve different traits. Convergent evolution is similar to parallel evolution, which occurs when two independent species evolve in the same direction and thus independently acquire similar characteristics; for instance, gliding frogs have evolved in parallel from multiple types of tree frog.

Many instances of convergent evolution are known in plants, including the repeated development of C4 photosynthesis, seed dispersal by fleshy fruits adapted to be eaten by animals, and carnivory.

Overview

Analogous & Homologous Structures
Homology and analogy in mammals and insects: on the horizontal axis, the structures are homologous in morphology, but different in function due to differences in habitat. On the vertical axis, the structures are analogous in function due to similar lifestyles but anatomically different with different phylogeny.[a]

In morphology, analogous traits arise when different species live in similar ways and/or a similar environment, and so face the same environmental factors. When occupying similar ecological niches (that is, a distinctive way of life) similar problems can lead to similar solutions.[1][2][3] The British anatomist Richard Owen was the first to identify the fundamental difference between analogies and homologies.[4]

In biochemistry, physical and chemical constraints on mechanisms have caused some active site arrangements such as the catalytic triad to evolve independently in separate enzyme superfamilies.[5]

In his 1989 book Wonderful Life, Stephen Jay Gould argued that if one could "rewind the tape of life [and] the same conditions were encountered again, evolution could take a very different course".[6] Simon Conway Morris disputes this conclusion, arguing that convergence is a dominant force in evolution, and given that the same environmental and physical constraints are at work, life will inevitably evolve toward an "optimum" body plan, and at some point, evolution is bound to stumble upon intelligence, a trait presently identified with at least primates, corvids, and cetaceans.[7]

Distinctions

Cladistics

In cladistics, a homoplasy is a trait shared by two or more taxa for any reason other than that they share a common ancestry. Taxa which do share ancestry are part of the same clade; cladistics seeks to arrange them according to their degree of relatedness to describe their phylogeny. Homoplastic traits caused by convergence are therefore, from the point of view of cladistics, confounding factors which could lead to an incorrect analysis.[8][9][10][11]

Atavism

In some cases, it is difficult to tell whether a trait has been lost and then re-evolved convergently, or whether a gene has simply been switched off and then re-enabled later. Such a re-emerged trait is called an atavism. From a mathematical standpoint, an unused gene (selectively neutral) has a steadily decreasing probability of retaining potential functionality over time. The time scale of this process varies greatly in different phylogenies; in mammals and birds, there is a reasonable probability of remaining in the genome in a potentially functional state for around 6 million years.[12]

Parallel vs. convergent evolution

Evolutionary trends
Evolution at an amino acid position. In each case, the left-hand species changes from having alanine (A) at a specific position in a protein in a hypothetical ancestor, and now has serine (S) there. The right-hand species may undergo divergent, parallel, or convergent evolution at this amino acid position relative to the first species.

When two species are similar in a particular character, evolution is defined as parallel if the ancestors were also similar, and convergent if they were not.[b] Some scientists have argued that there is a continuum between parallel and convergent evolution, while others maintain that despite some overlap, there are still important distinctions between the two.[13][14][15]

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, the striking example of similar placental and marsupial forms is described by Richard Dawkins in The Blind Watchmaker as a case of convergent evolution, because mammals on each continent had a long evolutionary history prior to the extinction of the dinosaurs under which to accumulate relevant differences.[16]

At molecular level

Triad convergence ser cys
Evolutionary convergence of serine and cysteine protease towards the same catalytic triads organisation of acid-base-nucleophile in different protease superfamilies. Shown are the triads of subtilisin, prolyl oligopeptidase, TEV protease, and papain.

Protease active sites

The enzymology of proteases provides some of the clearest examples of convergent evolution. These examples reflect the intrinsic chemical constraints on enzymes, leading evolution to converge on equivalent solutions independently and repeatedly.[5][17]

Serine and cysteine proteases use different amino acid functional groups (alcohol or thiol) as a nucleophile. In order to activate that nucleophile, they orient an acidic and a basic residue in a catalytic triad. The chemical and physical constraints on enzyme catalysis have caused identical triad arrangements to evolve independently more than 20 times in different enzyme superfamilies.[5]

Threonine proteases use the amino acid threonine as their catalytic nucleophile. Unlike cysteine and serine, threonine is a secondary alcohol (i.e. has a methyl group). The methyl group of threonine greatly restricts the possible orientations of triad and substrate, as the methyl clashes with either the enzyme backbone or the histidine base. Consequently, most threonine proteases use an N-terminal threonine in order to avoid such steric clashes. Several evolutionarily independent enzyme superfamilies with different protein folds use the N-terminal residue as a nucleophile. This commonality of active site but difference of protein fold indicates that the active site evolved convergently in those families.[5][18]

Nucleic acids

Convergence occurs at the level of DNA and the amino acid sequences produced by translating structural genes into proteins. Studies have found convergence in amino acid sequences in echolocating bats and the dolphin;[19] among marine mammals;[20] between giant and red pandas;[21] and between the thylacine and canids.[22] Convergence has also been detected in a type of non-coding DNA, cis-regulatory elements, such as in their rates of evolution; this could indicate either positive selection or relaxed purifying selection.[23]

In animal morphology

Ichthyosaur vs dolphin
Dolphins and ichthyosaurs converged on many adaptations for fast swimming.

Bodyplans

Swimming animals including fish such as herrings, marine mammals such as dolphins, and ichthyosaurs (of the Mesozoic) all converged on the same streamlined shape.[24][25] The fusiform bodyshape (a tube tapered at both ends) adopted by many aquatic animals is an adaptation to enable them to travel at high speed in a high drag environment.[26] Similar body shapes are found in the earless seals and the eared seals: they still have four legs, but these are strongly modified for swimming.[27]

The marsupial fauna of Australia and the placental mammals of the Old World have several strikingly similar forms, developed in two clades, isolated from each other.[7] The body and especially the skull shape of the thylacine (Tasmanian wolf) converged with those of Canidae such as the red fox, Vulpes vulpes.[28]

Vulpes vulpes skeleton

Red fox skeleton

Beutelwolf fg01

Skulls of thylacine (left), timber wolf (right)

Beutelwolfskelett brehm

Thylacine skeleton

Echolocation

As a sensory adaptation, echolocation has evolved separately in cetaceans (dolphins and whales) and bats, but from the same genetic mutations.[29][30]

Eyes

Evolution eye
The camera eyes of vertebrates (left) and cephalopods (right) developed independently and are wired differently; for instance, optic nerve fibres reach the vertebrate retina from the front, creating a blind spot.[31]

One of the best-known examples of convergent evolution is the camera eye of cephalopods (such as squid and octopus), vertebrates (including mammals) and cnidaria (such as jellyfish).[32] Their last common ancestor had at most a simple photoreceptive spot, but a range of processes led to the progressive refinement of camera eyes — with one sharp difference: the cephalopod eye is "wired" in the opposite direction, with blood and nerve vessels entering from the back of the retina, rather than the front as in vertebrates. As a result, cephalopods lack a blind spot.[7]

Flight

Homology
Vertebrate wings are partly homologous (from forelimbs), but analogous as organs of flight in (1) pterosaurs, (2) bats, (3) birds, evolved separately.

Birds and bats have homologous limbs because they are both ultimately derived from terrestrial tetrapods, but their flight mechanisms are only analogous, so their wings are examples of functional convergence. The two groups have powered flight, evolved independently. Their wings differ substantially in construction. The bat wing is a membrane stretched across four extremely elongated fingers and the legs. The airfoil of the bird wing is made of feathers, strongly attached to the forearm (the ulna) and the highly fused bones of the wrist and hand (the carpometacarpus), with only tiny remnants of two fingers remaining, each anchoring a single feather. So, while the wings of bats and birds are functionally convergent, they are not anatomically convergent.[3][33] Birds and bats also share a high concentration of cerebrosides in the skin of their wings. This improves skin flexibility, a trait useful for flying animals; other mammals have a far lower concentration.[34] The extinct pterosaurs independently evolved wings from their fore- and hindlimbs, while insects have wings that evolved separately from different organs.[35]

Flying squirrels and sugar gliders are much alike in their body plans, with gliding wings stretched between their limbs, but flying squirrels are placental mammals while sugar gliders are marsupials, widely separated within the mammal lineage.[36]

Insect mouthparts

Insect mouthparts show many examples of convergent evolution. The mouthparts of different insect groups consist of a set of homologous organs, specialised for the dietary intake of that insect group. Convergent evolution of many groups of insects led from original biting-chewing mouthparts to different, more specialised, derived function types. These include, for example, the proboscis of flower-visiting insects such as bees and flower beetles,[37][38][39] or the biting-sucking mouthparts of blood-sucking insects such as fleas and mosquitos.

Opposable thumbs

Opposable thumbs allowing the grasping of objects are most often associated with primates, like humans, monkeys, apes, and lemurs. Opposable thumbs also evolved in giant pandas, but these are completely different in structure, having six fingers including the thumb, which develops from a wrist bone entirely separately from other fingers.[40]

Primates

Veronika Loncká
Angolan women
(미쓰와이프) 제작기영상 엄정화 3m3s
Convergent evolution human skin color map
Despite the similar lightening of skin colour after moving out of Africa, different genes were involved in European (left) and East-Asian (right) lineages.

Convergent evolution in humans includes blue eye colour and light skin colour. When humans migrated out of Africa, they moved to more northern latitudes with less intense sunlight. It was beneficial to them to reduce their skin pigmentation. It appears certain that there was some lightening of skin colour before European and East Asian lineages diverged, as there are some skin-lightening genetic differences that are common to both groups. However, after the lineages diverged and became genetically isolated, the skin of both groups lightened more, and that additional lightening was due to different genetic changes.[41]

Humans Lemurs
A blue eye
Eye See You (2346693372)
Eulemur mongoz (male - face)
Blue-eyed black lemur
Despite the similarity of appearance, the genetic basis of blue eyes is different in humans and lemurs.

Lemurs and humans are both primates. Ancestral primates had brown eyes, as most primates do today. The genetic basis of blue eyes in humans has been studied in detail and much is known about it. It is not the case that one gene locus is responsible, say with brown dominant to blue eye colour. However, a single locus is responsible for about 80% of the variation. In lemurs, the differences between blue and brown eyes are not completely known, but the same gene locus is not involved.[42]

In plants

Chelidonium majus seeds
In myrmecochory, seeds such as those of Chelidonium majus have a hard coating and an attached oil body, an elaiosome, for dispersal by ants.

Carbon fixation

While convergent evolution is often illustrated with animal examples, it has often occurred in plant evolution. For instance, C4 photosynthesis, one of the three major carbon-fixing biochemical processes, has arisen independently up to 40 times.[43][44] About 7,600 plant species of angiosperms use C4 carbon fixation, with many monocots including 46% of grasses such as maize and sugar cane,[45][46] and dicots including several species in the Chenopodiaceae and the Amaranthaceae.[47][48]

Fruits

A good example of convergence in plants is the evolution of edible fruits such as apples. These pomes incorporate (five) carpels and their accessory tissues forming the apple's core, surrounded by structures from outside the botanical fruit, the receptacle or hypanthium. Other edible fruits include other plant tissues;[49] for example, the fleshy part of a tomato is the walls of the pericarp.[50] This implies convergent evolution under selective pressure, in this case the competition for seed dispersal by animals through consumption of fleshy fruits.[51]

Seed dispersal by ants (myrmecochory) has evolved independently more than 100 times, and is present in more than 11,000 plant species. It is one of the most dramatic examples of convergent evolution in biology.[52]

Carnivory

Chitinase4TC
Molecular convergence in carnivorous plants

Carnivory has evolved multiple times independently in plants in widely separated groups. In three species studied, Cephalotus follicularis, Nepenthes alata and Sarracenia purpurea, there has been convergence at the molecular level. Carnivorous plants secrete enzymes into the digestive fluid they produce. By studying phosphatase, glycoside hydrolase, glucanase, RNAse and chitinase enzymes as well as a pathogenesis-related protein and a thaumatin-related protein, the authors found many convergent amino acid substitutions. These changes were not at the enzymes' catalytic sites, but rather on the exposed surfaces of the proteins, where they might interact with other components of the cell or the digestive fluid. The authors also found that homologous genes in the non-carnivorous plant Arabidopsis thaliana tend to have their expression increased when the plant is stressed, leading the authors to suggest that stress-responsive proteins have often been co-opted[c] in the repeated evolution of carnivory.[53]

Methods of inference

Phenotypic-landscape-inference-reveals-multiple-evolutionary-paths-toC4-photosynthesis-elife00961fs002
Angiosperm phylogeny of orders based on classification by the Angiosperm Phylogeny Group. The figure shows the number of inferred independent origins of C3-C4 photosynthesis and C4 photosynthesis in parentheses.

Phylogenetic reconstruction and ancestral state reconstruction proceed by assuming that evolution has occurred without convergence. Convergent patterns may, however, appear at higher levels in a phylogenetic reconstruction, and are sometimes explicitly sought by investigators. The methods applied to infer convergent evolution depend on whether pattern-based or process-based convergence is expected. Pattern-based convergence is the broader term, for when two or more lineages independently evolve patterns of similar traits. Process-based convergence is when the convergence is due to similar forces of natural selection.[54]

Pattern-based measures

Earlier methods for measuring convergence incorporate ratios of phenotypic and phylogenetic distance by simulating evolution with a Brownian motion model of trait evolution along a phylogeny.[55][56] More recent methods also quantify the strength of convergence.[57] One drawback to keep in mind is that these methods can confuse long-term stasis with convergence due to phenotypic similarities. Stasis occurs when there is little evolutionary change among taxa.[54]

Distance-based measures assess the degree of similarity between lineages over time. Frequency-based measures assess the number of lineages that have evolved in a particular trait space.[54]

Process-based measures

Methods to infer process-based convergence fit models of selection to a phylogeny and continuous trait data to determine whether the same selective forces have acted upon lineages. This uses the Ornstein-Uhlenbeck (OU) process to test different scenarios of selection. Other methods rely on an a priori specification of where shifts in selection have occurred.[58]

See also

  • Incomplete lineage sorting: the presence of multiple alleles in ancestral populations might lead to the impression that convergent evolution has occurred.

Notes

  1. ^ However, evolutionary developmental biology has identified deep homology between insect and mammal body plans, to the surprise of many biologists.
  2. ^ However, all organisms share a common ancestor more or less recently, so the question of how far back to look in evolutionary time and how similar the ancestors need to be for one to consider parallel evolution to have taken place is not entirely resolved within evolutionary biology.
  3. ^ The prior existence of suitable structures has been called pre-adaptation or exaptation.

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Birds of Australia

Australia and its offshore islands and territories have 898 recorded bird species as of 2014. Of the recorded birds, 165 are considered vagrant or accidental visitors, of the remainder over 45% are classified as Australian endemics: found nowhere else on earth. It has been suggested that up to 10% of Australian bird species may go extinct by the year 2100 as a result of climate change.Australian species range from the tiny 8 cm weebill to the huge, flightless emu. Many species of Australian birds will immediately seem familiar to visitors from the Northern Hemisphere - Australian wrens look and act much like northern wrens and Australian robins seem to be close relatives of the northern robins, but in fact the majority of Australian passerines are descended from the ancestors of the crow family, and the close resemblance is misleading: the cause is not genetic relatedness but convergent evolution.

For example, almost any land habitat offers a nice home for a small bird that specialises in finding small insects: the form best fitted to that task is one with long legs for agility and obstacle clearance, moderately-sized wings optimised for quick, short flights, and a large, upright tail for rapid changes of direction. In consequence, the unrelated birds that fill that role in the Americas and in Australia look and act as though they are close relatives.

Australian birds which show convergent evolution with Northern Hemisphere species:

honeyeaters (resemble sunbirds)

sittellas (resemble nuthatches)

Australasian babblers (resemble scimitar babblers)

Australian robins (resemble Old World chats)

Scrub robins (resemble thrushes)

Catkin

A catkin or ament is a slim, cylindrical flower cluster (a spike), with inconspicuous or no petals, usually wind-pollinated (anemophilous) but sometimes insect-pollinated (as in Salix). They contain many, usually unisexual flowers, arranged closely along a central stem which is often drooping. They are found in many plant families, including Betulaceae, Fagaceae, Moraceae, and Salicaceae. For some time, they were believed to be a key synapomorphy among the proposed Hamamelididae, also known as Amentiferae (i.e., literally plants bearing aments). Based on molecular phylogeny work, it is now believed that Hamamelididae is a polyphyletic group. This suggests that the catkin flower arrangement has arisen at least twice independently by convergent evolution, in Fagales and in Salicaceae. Such a convergent evolution raises questions about what the ancestral inflorescence characters might be and how catkins did evolve in these two lineages.

In many of these plants, only the male flowers form catkins, and the female flowers are single (hazel, oak), a cone (alder) or other types (mulberry). In other plants (such as poplar) both male and female flowers are borne in catkins.

Catkin-bearing plants include many other trees or shrubs such as birch, willow, hickory, sweet chestnut and sweetfern (Comptonia).

The word catkin is a loanword from the old Dutch katteken, meaning "kitten", on account of the resemblance to a kitten's tail. Ament is from the Latin amentum, meaning "thong" or "strap".

Cephalopod eye

Cephalopods, as active marine predators, possess sensory organs specialized for use in aquatic conditions. They have a camera-type eye which consists of an iris, a circular lens, vitreous cavity (eye gel), pigment cells, and photoreceptor cells that translate light from the light-sensitive retina into nerve signals which travel along the optic nerve to the brain. For the past 140 years, the camera-type cephalopod eye has been compared with the vertebrate eye as an example of convergent evolution, where both types of organisms have independently evolved the camera-eye trait and both share similar functionality. Contention exists on whether this is truly convergent evolution or parallel evolution. Unlike the vertebrate camera eye, the cephalopods' form as invaginations of the body surface (rather than outgrowths of the brain), and consequently they lack a cornea. Unlike the vertebrate eye, a cephalopod eye is focused through movement, much like the lens of a camera or telescope, rather than changing shape as the lens in the human eye does. The eye is approximately spherical, as is the lens, which is fully internal.Cephalopods' eyes develop in such a way that they have retinal axons that pass over the back of the retina, so the optic nerve does not have to pass through the photoreceptor layer to exit the eye and do not have the natural, central, physiological blind spot of vertebrates.The crystalins used in the lens appear to have developed independently from vertebrate crystalins, suggesting a homoplasious origin of the lens.Most cephalopods possess complex extraocular muscle systems that allow for very fine control over the gross positioning of the eyes. Octopuses possess an autonomic response that maintains the orientation of their pupils such that they are always horizontal.

Cleaner fish

Cleaner fish are fish that provide a service to other species by removing dead skin and ectoparasites. Although the animal being cleaned typically is another fish, it can also involve aquatic reptiles (sea turtles and marine iguana), mammals (manatees and whales) or octopuses. The cleaning symbiosis is an example of mutualism, an ecological interaction that benefits both parties involved. However, the cleaner fish may sometimes cheat and consume mucus or tissue, thus creating a form of parasitism. A wide variety of fish including wrasse, cichlids, catfish, pipefish, and gobies display cleaning behaviors. Similar behavior is found in other groups of animals, such as cleaner shrimps.

Cleaner fish advertise their services with conspicuous coloration, often displaying a brilliant blue stripe that spans the length of the body. This adaptation has evolved independently in different species of cleaner fish, making it an example of convergent evolution. Other species of fish, called mimics, imitate the behavior and phenotype of cleaner fish to gain access to client fish tissue. This is another example of convergent evolution.

Corticioid fungi

The corticioid fungi are a group of fungi in the Basidiomycota typically having effused, smooth basidiocarps (fruit bodies) that are formed on the undersides of dead tree trunks or branches. They are sometimes colloquially called crust fungi or patch fungi. Originally such fungi were referred to the genus Corticium ("corticioid" means Corticium-like) and subsequently to the family Corticiaceae, but it is now known that all corticioid species are not necessarily closely related. The fact that they look similar is an example of convergent evolution. Since they are often studied as a group, it is convenient to retain the informal (non-taxonomic) name of "corticioid fungi" and this term is frequently used in research papers and other texts.

Divergent evolution

Divergent evolution or divergent selection is the accumulation of differences between closely related species populations, leading to speciation. Divergent evolution is typically exhibited when two populations become separated by a geographic barrier (such as in allopatric or peripatric speciation) and experience different selective pressures that drive adaptations to their new environment. After many generations and continual evolution, the populations become unable to interbreed with one another. The American naturalist J. T. Gulick (1832-1923) was the first to use the term "divergent evolution", with its use becoming widespread in modern evolutionary literature. Classic examples of divergence in nature are the adaptive radiation of the finches of the Galapagos or the coloration differences in populations of a species that live in different habitats such as with pocket mice and fence lizards.The term can also be applied in molecular evolution, such as to proteins that derive from homologous genes. Both orthologous genes (resulting from a speciation event) and paralogous genes (resulting from gene duplication) can illustrate divergent evolution. Through gene duplication, it is possible for divergent evolution to occur between two genes within a species. Similarities between species that have diverged are due to their common origin, so such similarities are homologies. In contrast, convergent evolution arises when an adaptation has arisen independently, creating analogous structures such as the wings of birds and of insects.

Elaiosome

Elaiosomes (Greek élaion "oil" and sóma "body") are fleshy structures that are attached to the seeds of many plant species. The elaiosome is rich in lipids and proteins, and may be variously shaped. Many plants have elaiosomes that attract ants, which take the seed to their nest and feed the elaiosome to their larvae. After the larvae have consumed the elaiosome, the ants take the seed to their waste disposal area, which is rich in nutrients from the ant frass and dead bodies, where the seeds germinate. This type of seed dispersal is termed myrmecochory from the Greek "ant" (myrmex) and "circular dance" (khoreíā). This type of symbiotic relationship appears to be mutualistic, more specifically dispersive mutualism according to Ricklefs, R.E. (2001), as the plant benefits because its seeds are dispersed to favorable germination sites, and also because it is planted (carried underground) by the ants.

Elaiosomes develop in various ways either from seed tissues (chalaza, funiculus, hilum, raphe-antiraphe) or from fruit tissues (exocarp, receptacle, flower tube, perigonium, style or spicule). The various origins and developmental pathways apparently all serve the same main function, i.e. attracting ants. Because elaiosomes are present in at least 11,000, but possibly up to 23,000 species of plants, elaiosomes are a dramatic example of convergent evolution in flowering plants.

Enzyme Commission number

The Enzyme Commission number (EC number) is a numerical classification scheme for enzymes, based on the chemical reactions they catalyze.

As a system of enzyme nomenclature, every EC number is associated with a recommended name for the respective enzyme.

Strictly speaking, EC numbers do not specify enzymes, but enzyme-catalyzed reactions. If different enzymes (for instance from different organisms) catalyze the same reaction, then they receive the same EC number. Furthermore, through convergent evolution, completely different protein folds can catalyze an identical reaction and therefore would be assigned an identical EC number (these are called non-homologous isofunctional enzymes, or NISE). By contrast, UniProt identifiers uniquely specify a protein by its amino acid sequence.

Euphorbiaceae

The Euphorbiaceae, the spurge family, is a large family of flowering plants. In common English, they are sometimes called euphorbias, which is also the name of a genus in the family. Most spurges such as Euphorbia paralias are herbs, but some, especially in the tropics, are shrubs or trees, such as Hevea brasiliensis. Some, such as Euphorbia canariensis, are succulent and resemble cacti because of convergent evolution. This family occurs mainly in the tropics, with the majority of the species in the Indo-Malayan region and tropical America a strong second. A large variety occurs in tropical Africa, but they are not as abundant or varied as in the two other tropical regions. However, Euphorbiaceae also has many species in nontropical areas such as the Mediterranean Basin, the Middle East, South Africa, and the southern United States.

Honeyeater

The honeyeaters are a large and diverse family, Meliphagidae, of small to medium-sized birds. The family includes the Australian chats, myzomelas, friarbirds, wattlebirds, miners and melidectes. They are most common in Australia and New Guinea, but also found in New Zealand, the Pacific islands as far east as Samoa and Tonga, and the islands to the north and west of New Guinea known as Wallacea. Bali, on the other side of the Wallace Line, has a single species.In total there are 187 species in 50 genera, roughly half of them native to Australia, many of the remainder occupying New Guinea. With their closest relatives, the Maluridae (Australian fairy-wrens), Pardalotidae (pardalotes), and Acanthizidae (thornbills, Australian warblers, scrubwrens, etc.), they comprise the superfamily Meliphagoidea and originated early in the evolutionary history of the oscine passerine radiation. Although honeyeaters look and behave very much like other nectar-feeding passerines around the world (such as the sunbirds and flowerpeckers), they are unrelated, and the similarities are the consequence of convergent evolution.

The extent of the evolutionary partnership between honeyeaters and Australasian flowering plants is unknown, but probably substantial. A great many Australian plants are fertilised by honeyeaters, particularly the Proteaceae, Myrtaceae, and Epacridaceae. It is known that the honeyeaters are important in New Zealand as well, and assumed that the same applies in other areas.

Humanoid

A humanoid (; from English human and -oid "resembling") is something that has an appearance resembling a human without actually being one. The earliest recorded use of the term, in 1870, referred to indigenous peoples in areas colonized by Europeans. By the 20th century, the term came to describe fossils which were morphologically similar, but not identical, to those of the human skeleton.Although this usage was common in the sciences for much of the 20th century, it is now considered rare. More generally, the term can refer to anything with distinctly human characteristics or adaptations, such as possessing opposable anterior forelimb-appendages (i.e. thumbs), visible spectrum-binocular vision (i.e. having two eyes), or biomechanic plantigrade-bipedalism (i.e. the ability to walk on heels and metatarsals in an upright position). Science fiction media frequently present sentient extraterrestrial lifeforms as humanoid as a byproduct of convergent evolution theory.

List of examples of convergent evolution

Convergent evolution — the repeated evolution of similar traits in multiple lineages which all ancestrally lack the trait — is rife in nature, as illustrated by the examples below. The ultimate cause of convergence is usually a similar evolutionary biome, as similar environments will select for similar traits in any species occupying the same ecological niche, even if those species are only distantly related. In the case of cryptic species, it can create species which are only distinguishable by analysing their genetics. Unrelated organisms often develop analogous structures by adapting to similar environments.

Mohoidae

Mohoidae is a family of Hawaiian species of recently extinct, nectarivorous songbirds in the genera Moho (ʻōʻō) and Chaetoptila (kioea). These now extinct birds form their own family, representing the only complete extinction of an entire avian family in modern times, when the disputed family Turnagridae is disregarded for being invalid.

Until recently, these birds were thought to belong to the family Meliphagidae (honeyeaters) due to their very similar appearance and behavior, including many morphological details. However, a 2008 study argued, on the basis of a phylogenetic analysis of DNA from museum specimens, that the genera Moho and Chaetoptila do not belong to the Meliphagidae but instead belong to a group that includes the waxwings and the palmchat; they appear especially close to the silky-flycatchers. Hawaiian honeyeaters did not evolve from the similar looking Australasian honeyeaters, but instead represent a striking case of convergent evolution. The authors proposed a family, Mohoidae, for these two extinct genera.

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.

Pill millipede

Pill millipedes are any members of two living (and one extinct) orders of millipedes, often grouped together into a single superorder, Oniscomorpha. The name Oniscomorpha refers to the millipedes' resemblance to certain woodlice (Oniscidea), also called pillbugs or "roly-polies". However, millipedes and woodlice are not closely related (belonging to the subphyla Myriapoda and Crustacea, respectively); rather, this is a case of convergent evolution.

Polyphyly

A polyphyletic group is a set of organisms, or other evolving elements, that have been grouped together but do not share an immediate common ancestor. The term is often applied to groups that share characteristics that appear to be similar but have not been inherited from common ancestors; these characteristics are known as homoplasies, and the development and phenomenon of homoplasies is known as convergent evolution. The arrangement of the members of a polyphyletic group is called a polyphyly.

Alternatively, polyphyletic is simply used to describe a group whose members come from multiple ancestral sources, regardless of similarity of characteristics. For example, the biological characteristic of warm-bloodedness evolved separately in the ancestors of mammals and the ancestors of birds. Other polyphyletic groups are for example algae, C4 photosynthetic plants, and edentates.Many biologists aim to avoid homoplasies in grouping taxa together and therefore it is frequently a goal to eliminate groups that are found to be polyphyletic. This is often the stimulus for major revisions of the classification schemes.

Researchers concerned more with ecology than with systematics may take polyphyletic groups as legitimate subject matter; the similarities in activity within the fungus group Alternaria, for example, can lead researchers to regard the group as a valid genus while acknowledging its polyphyly.

Proleg

A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the order Lepidoptera, though they can also be found on other larval insects such as sawflies and a few types of flies. In all the orders in which they appear, mainly Hymenoptera and Lepidoptera, prolegs of any form evolved independently of each other by convergent evolution.Prolegs of lepidopteran larvae have a small circle of gripping hooks, called "crochets". The arrangement of the crochets can be helpful in identification to family level. Although the point has been debated, prolegs are not widely regarded as true legs, derived from the primitive uniramous limbs. Certainly in their morphology they are not jointed, and so lack the five segments (coxa, trochanter, femur, tibia, tarsus) of thoracic insect legs. Prolegs do have limited musculature, but much of their movement is hydraulically powered.

Tibiotarsus

The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia.

A similar structure also occurred in the Mesozoic Heterodontosauridae. These small ornithischian dinosaurs were unrelated to birds and the similarity of their foot bones is best explained by convergent evolution.

Tree frog

A tree frog is any species of frog that spends a major portion of its lifespan in trees, known as an arboreal state. Several lineages of frogs among the Neobatrachia have given rise to tree frogs, although they are not closely related to each other.

Many millions of years of convergent evolution have resulted in almost identical morphology and ecologies. They are so similar as regards their ecological niche that in one biome where one group of tree frogs occurs, the other is almost always absent. The last common ancestor of some such tree frog groups lived long before the extinction of the dinosaurs.

Patterns of evolution
Signals

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