123-425: Beewolves (genus Philanthus ), also known as bee-hunters or bee-killer wasps , are solitary, predatory wasps , most of which prey on bees , hence their common name. The adult females dig tunnels in the ground for nesting, while the territorial males mark twigs and other objects with pheromones to claim the territory from competing males. As with all other apoid wasps, the larvae are carnivorous , forcing
246-455: A mutation (the deletion of two nucleotides ) that inactives it. These changes are explained by the fact that its prey does not need to be subdued. Several groups of predatory fish have the ability to detect, track, and sometimes, as in the electric ray , to incapacitate their prey by sensing and generating electric fields . The electric organ is derived from modified nerve or muscle tissue. Physiological adaptations to predation include
369-487: A DNA sequence within a chromosome is known as a locus . If the DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles. DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene, the new allele may affect the trait that the gene controls, altering the phenotype of the organism. However, while this simple correspondence between an allele and
492-570: A GC-biased E. coli mutator strain in 1967, along with the proposal of the neutral theory , established the plausibility of mutational explanations for molecular patterns, which are now common in the molecular evolution literature. For instance, mutation biases are frequently invoked in models of codon usage. Such models also include effects of selection, following the mutation-selection-drift model, which allows both for mutation biases and differential selection based on effects on translation. Hypotheses of mutation bias have played an important role in
615-418: A better choice. If it chooses pursuit, its physical capabilities determine the mode of pursuit (e.g., ambush or chase). Having captured the prey, it may also need to expend energy handling it (e.g., killing it, removing any shell or spines, and ingesting it). Predators have a choice of search modes ranging from sit-and-wait to active or widely foraging . The sit-and-wait method is most suitable if
738-851: A broad range of taxa including arthropods. They are common among insects, including mantids, dragonflies , lacewings and scorpionflies . In some species such as the alderfly , only the larvae are predatory (the adults do not eat). Spiders are predatory, as well as other terrestrial invertebrates such as scorpions ; centipedes ; some mites , snails and slugs ; nematodes ; and planarian worms . In marine environments, most cnidarians (e.g., jellyfish , hydroids ), ctenophora (comb jellies), echinoderms (e.g., sea stars , sea urchins , sand dollars , and sea cucumbers ) and flatworms are predatory. Among crustaceans , lobsters , crabs , shrimps and barnacles are predators, and in turn crustaceans are preyed on by nearly all cephalopods (including octopuses , squid and cuttlefish ). Seed predation
861-514: A chromosome becoming duplicated (usually by genetic recombination ), which can introduce extra copies of a gene into a genome. Extra copies of genes are a major source of the raw material needed for new genes to evolve. This is important because most new genes evolve within gene families from pre-existing genes that share common ancestors. For example, the human eye uses four genes to make structures that sense light: three for colour vision and one for night vision ; all four are descended from
984-582: A food trap, mechanical stimulation, and electrical impulses to eventually catch and consume its prey. Some carnivorous fungi catch nematodes using either active traps in the form of constricting rings, or passive traps with adhesive structures. Many species of protozoa ( eukaryotes ) and bacteria ( prokaryotes ) prey on other microorganisms; the feeding mode is evidently ancient, and evolved many times in both groups. Among freshwater and marine zooplankton , whether single-celled or multi-cellular, predatory grazing on phytoplankton and smaller zooplankton
1107-471: A huge gulp of water and filtering it through their feathery baleen plates. Pursuit predators may be social , like the lion and wolf that hunt in groups, or solitary. Once the predator has captured the prey, it has to handle it: very carefully if the prey is dangerous to eat, such as if it possesses sharp or poisonous spines, as in many prey fish. Some catfish such as the Ictaluridae have spines on
1230-725: A kill, and the coyote can be either solitary or social. Other solitary predators include the northern pike, wolf spiders and all the thousands of species of solitary wasps among arthropods, and many microorganisms and zooplankton . Under the pressure of natural selection , predators have evolved a variety of physical adaptations for detecting, catching, killing, and digesting prey. These include speed, agility, stealth, sharp senses, claws, teeth, filters, and suitable digestive systems. For detecting prey , predators have well-developed vision , smell , or hearing . Predators as diverse as owls and jumping spiders have forward-facing eyes, providing accurate binocular vision over
1353-539: A long distance, sometimes for hours at a time. The method is used by human hunter-gatherers and by canids such as African wild dogs and domestic hounds. The African wild dog is an extreme persistence predator, tiring out individual prey by following them for many miles at relatively low speed. A specialised form of pursuit predation is the lunge feeding of baleen whales . These very large marine predators feed on plankton , especially krill , diving and actively swimming into concentrations of plankton, and then taking
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#17330846326571476-402: A manner similar to a sequence of letters spelling out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called genes; different genes have different sequences of bases. Within cells, each long strand of DNA is called a chromosome . The specific location of
1599-405: A nutrient in a long-term laboratory experiment , Flavobacterium evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing, and the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic pesticide pentachlorophenol . An interesting but still controversial idea is that some adaptations might increase
1722-445: A patch and decide whether to spend time searching for prey in it. This may involve some knowledge of the preferences of the prey; for example, ladybirds can choose a patch of vegetation suitable for their aphid prey. To capture prey, predators have a spectrum of pursuit modes that range from overt chase ( pursuit predation ) to a sudden strike on nearby prey ( ambush predation ). Another strategy in between ambush and pursuit
1845-611: A phenomenon termed de novo gene birth . The generation of new genes can also involve small parts of several genes being duplicated, with these fragments then recombining to form new combinations with new functions ( exon shuffling ). When new genes are assembled from shuffling pre-existing parts, domains act as modules with simple independent functions, which can be mixed together to produce new combinations with new and complex functions. For example, polyketide synthases are large enzymes that make antibiotics ; they contain up to 100 independent domains that each catalyse one step in
1968-535: A population are therefore more likely to be replaced by the offspring of parents with favourable characteristics for that environment. In the early 20th century, competing ideas of evolution were refuted and evolution was combined with Mendelian inheritance and population genetics to give rise to modern evolutionary theory. In this synthesis the basis for heredity is in DNA molecules that pass information from generation to generation. The processes that change DNA in
2091-575: A population include natural selection, genetic drift, mutation , and gene flow . All life on Earth—including humanity —shares a last universal common ancestor (LUCA), which lived approximately 3.5–3.8 billion years ago. The fossil record includes a progression from early biogenic graphite to microbial mat fossils to fossilised multicellular organisms . Existing patterns of biodiversity have been shaped by repeated formations of new species ( speciation ), changes within species ( anagenesis ), and loss of species ( extinction ) throughout
2214-430: A population is not critical, but instead a measure known as the effective population size. The effective population is usually smaller than the total population since it takes into account factors such as the level of inbreeding and the stage of the lifecycle in which the population is the smallest. The effective population size may not be the same for every gene in the same population. It is usually difficult to measure
2337-433: A population through neutral transitions due to the principles of excess capacity, presuppression, and ratcheting, and it has been applied in areas ranging from the origins of the spliceosome to the complex interdependence of microbial communities . The time it takes a neutral allele to become fixed by genetic drift depends on population size; fixation is more rapid in smaller populations. The number of individuals in
2460-425: A powerful selective effect on prey, and the prey develop antipredator adaptations such as warning coloration , alarm calls and other signals , camouflage , mimicry of well-defended species, and defensive spines and chemicals. Sometimes predator and prey find themselves in an evolutionary arms race , a cycle of adaptations and counter-adaptations. Predation has been a major driver of evolution since at least
2583-785: A predator, while small prey might prove hard to find and in any case provide less of a reward. This has led to a correlation between the size of predators and their prey. Size may also act as a refuge for large prey. For example, adult elephants are relatively safe from predation by lions, but juveniles are vulnerable. Members of the cat family such as the snow leopard (treeless highlands), tiger (grassy plains, reed swamps), ocelot (forest), fishing cat (waterside thickets), and lion (open plains) are camouflaged with coloration and disruptive patterns suiting their habitats. In aggressive mimicry , certain predators, including insects and fishes, make use of coloration and behaviour to attract prey. Female Photuris fireflies , for example, copy
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#17330846326572706-421: A preferred target is scarce. When prey have a clumped (uneven) distribution, the optimal strategy for the predator is predicted to be more specialized as the prey are more conspicuous and can be found more quickly; this appears to be correct for predators of immobile prey, but is doubtful with mobile prey. In size-selective predation, predators select prey of a certain size. Large prey may prove troublesome for
2829-499: A range of genes from bacteria, fungi and plants. Viruses can also carry DNA between organisms, allowing transfer of genes even across biological domains . Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and bacteria, during the acquisition of chloroplasts and mitochondria . It is possible that eukaryotes themselves originated from horizontal gene transfers between bacteria and archaea . Some heritable changes cannot be explained by changes to
2952-430: A reduction in scope when it was discovered that (1) GC-biased gene conversion makes an important contribution to composition in diploid organisms such as mammals and (2) bacterial genomes frequently have AT-biased mutation. Contemporary thinking about the role of mutation biases reflects a different theory from that of Haldane and Fisher. More recent work showed that the original "pressures" theory assumes that evolution
3075-820: A relatively narrow field of view, whereas prey animals often have less acute all-round vision. Animals such as foxes can smell their prey even when it is concealed under 2 feet (60 cm) of snow or earth. Many predators have acute hearing, and some such as echolocating bats hunt exclusively by active or passive use of sound. Predators including big cats , birds of prey , and ants share powerful jaws, sharp teeth, or claws which they use to seize and kill their prey. Some predators such as snakes and fish-eating birds like herons and cormorants swallow their prey whole; some snakes can unhinge their jaws to allow them to swallow large prey, while fish-eating birds have long spear-like beaks that they use to stab and grip fast-moving and slippery prey. Fish and other predators have developed
3198-478: A significant amount of energy, to locate each food patch. For example, the black-browed albatross regularly makes foraging flights to a range of around 700 kilometres (430 miles), up to a maximum foraging range of 3,000 kilometres (1,860 miles) for breeding birds gathering food for their young. With static prey, some predators can learn suitable patch locations and return to them at intervals to feed. The optimal foraging strategy for search has been modelled using
3321-452: A single ancestral gene. New genes can be generated from an ancestral gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has been duplicated because it increases the redundancy of the system; one gene in the pair can acquire a new function while the other copy continues to perform its original function. Other types of mutations can even generate entirely new genes from previously noncoding DNA,
3444-419: A single chromosome compared to expectations , which is called their linkage disequilibrium . A set of alleles that is usually inherited in a group is called a haplotype . This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the population; this effect
3567-406: A small animal, gulping the prey in an extremely rapid movement when it is within range. Many smaller predators such as the box jellyfish use venom to subdue their prey, and venom can also aid in digestion (as is the case for rattlesnakes and some spiders ). The marbled sea snake that has adapted to egg predation has atrophied venom glands, and the gene for its three finger toxin contains
3690-523: A smaller area. For example, when mixed flocks of birds forage, the birds in front flush out insects that are caught by the birds behind. Spinner dolphins form a circle around a school of fish and move inwards, concentrating the fish by a factor of 200. By hunting socially chimpanzees can catch colobus monkeys that would readily escape an individual hunter, while cooperating Harris hawks can trap rabbits. Predators of different species sometimes cooperate to catch prey. In coral reefs , when fish such as
3813-420: A species or population, in particular shifts in allele frequency and adaptation. Macroevolution is the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one—the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. For instance, a large amount of variation among individuals allows
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3936-544: A species to rapidly adapt to new habitats , lessening the chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it more likely that part of the population will become isolated. In this sense, microevolution and macroevolution might involve selection at different levels—with microevolution acting on genes and organisms, versus macroevolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction. A common misconception
4059-411: A striking example are people with the inherited trait of albinism , who do not tan at all and are very sensitive to sunburn . Heritable characteristics are passed from one generation to the next via DNA , a molecule that encodes genetic information. DNA is a long biopolymer composed of four types of bases. The sequence of bases along a particular DNA molecule specifies the genetic information, in
4182-472: A system in which organisms interact with every other element, physical as well as biological , in their local environment. Eugene Odum , a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and material cycles (i.e., exchange of materials between living and nonliving parts) within
4305-428: A trait works in some cases, most traits are influenced by multiple genes in a quantitative or epistatic manner. Evolution can occur if there is genetic variation within a population. Variation comes from mutations in the genome, reshuffling of genes through sexual reproduction and migration between populations ( gene flow ). Despite the constant introduction of new variation through mutation and gene flow, most of
4428-440: A variety of defences including the ability to hear the echolocation calls. Many pursuit predators that run on land, such as wolves, have evolved long limbs in response to the increased speed of their prey. Their adaptations have been characterized as an evolutionary arms race , an example of the coevolution of two species. In a gene centered view of evolution , the genes of predator and prey can be thought of as competing for
4551-481: A wide variety of bees such as the American bumblebee, Bombus pensylvanicus , or other hymenopterans , including conspecifics. They are notable for stinging their prey in a membranous location on the ventral surface, where the venom quickly paralyzes major voluntary muscles, yet does not kill the prey. The prey may attempt to sting in return, but it is always grabbed in such a way that only well-armored portions of
4674-477: Is ballistic interception , where a predator observes and predicts a prey's motion and then launches its attack accordingly. Ambush or sit-and-wait predators are carnivorous animals that capture prey by stealth or surprise. In animals, ambush predation is characterized by the predator's scanning the environment from a concealed position until a prey is spotted, and then rapidly executing a fixed surprise attack. Vertebrate ambush predators include frogs, fish such as
4797-454: Is directional selection , which is a shift in the average value of a trait over time—for example, organisms slowly getting taller. Secondly, disruptive selection is selection for extreme trait values and often results in two different values becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilising selection there
4920-442: Is more noticeable . Indeed, the evolution of microorganisms is particularly important to evolutionary research since their rapid reproduction allows the study of experimental evolution and the observation of evolution and adaptation in real time. Adaptation is the process that makes organisms better suited to their habitat. Also, the term adaptation may refer to a trait that is important for an organism's survival. For example,
5043-427: Is a byproduct of this process that may sometimes be adaptively beneficial. Gene flow is the exchange of genes between populations and between species. It can therefore be a source of variation that is new to a population or to a species. Gene flow can be caused by the movement of individuals between separate populations of organisms, as might be caused by the movement of mice between inland and coastal populations, or
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5166-417: Is an inherited characteristic and an individual might inherit the "brown-eye trait" from one of their parents. Inherited traits are controlled by genes and the complete set of genes within an organism's genome (genetic material) is called its genotype . The complete set of observable traits that make up the structure and behaviour of an organism is called its phenotype . Some of these traits come from
5289-464: Is based on standing variation: when evolution depends on events of mutation that introduce new alleles, mutational and developmental biases in the introduction of variation (arrival biases) can impose biases on evolution without requiring neutral evolution or high mutation rates. Several studies report that the mutations implicated in adaptation reflect common mutation biases though others dispute this interpretation. Recombination allows alleles on
5412-407: Is called deep homology . During evolution, some structures may lose their original function and become vestigial structures. Such structures may have little or no function in a current species, yet have a clear function in ancestral species, or other closely related species. Examples include pseudogenes , the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds,
5535-692: Is called genetic hitchhiking or genetic draft. Genetic draft caused by the fact that some neutral genes are genetically linked to others that are under selection can be partially captured by an appropriate effective population size. A special case of natural selection is sexual selection, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent among males of several animal species. Although sexually favoured, traits such as cumbersome antlers, mating calls, large body size and bright colours often attract predation, which compromises
5658-494: Is common, and found in many species of nanoflagellates , dinoflagellates , ciliates , rotifers , a diverse range of meroplankton animal larvae, and two groups of crustaceans, namely copepods and cladocerans . To feed, a predator must search for, pursue and kill its prey. These actions form a foraging cycle. The predator must decide where to look for prey based on its geographical distribution; and once it has located prey, it must assess whether to pursue it or to wait for
5781-402: Is distinct from scavenging on dead prey, though many predators also scavenge ; it overlaps with herbivory , as seed predators and destructive frugivores are predators. Predators may actively search for or pursue prey or wait for it, often concealed. When prey is detected, the predator assesses whether to attack it. This may involve ambush or pursuit predation , sometimes after stalking
5904-594: Is measured by an organism's ability to survive and reproduce, which determines the size of its genetic contribution to the next generation. However, fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes. For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make little genetic contribution to future generations and would thus have low fitness. If an allele increases fitness more than
6027-514: Is restricted to mammals, birds, and insects but is found in almost all terrestrial ecosystems. Egg predation includes both specialist egg predators such as some colubrid snakes and generalists such as foxes and badgers that opportunistically take eggs when they find them. Some plants, like the pitcher plant , the Venus fly trap and the sundew , are carnivorous and consume insects . Methods of predation by plants varies greatly but often involves
6150-421: Is selection against extreme trait values on both ends, which causes a decrease in variance around the average value and less diversity. This would, for example, cause organisms to eventually have a similar height. Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem , that is,
6273-430: Is size. Prey that is too small may not be worth the trouble for the amount of energy it provides. Too large, and it may be too difficult to capture. For example, a mantid captures prey with its forelegs and they are optimized for grabbing prey of a certain size. Mantids are reluctant to attack prey that is far from that size. There is a positive correlation between the size of a predator and its prey. A predator may assess
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#17330846326576396-429: Is that evolution has goals, long-term plans, or an innate tendency for "progress", as expressed in beliefs such as orthogenesis and evolutionism; realistically, however, evolution has no long-term goal and does not necessarily produce greater complexity. Although complex species have evolved, they occur as a side effect of the overall number of organisms increasing, and simple forms of life still remain more common in
6519-447: Is the nearly neutral theory , according to which a mutation that would be effectively neutral in a small population is not necessarily neutral in a large population. Other theories propose that genetic drift is dwarfed by other stochastic forces in evolution, such as genetic hitchhiking, also known as genetic draft. Another concept is constructive neutral evolution (CNE), which explains that complex systems can emerge and spread into
6642-500: 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 certain characteristics becoming more or less common within a population over successive generations. The process of evolution has given rise to biodiversity at every level of biological organisation . The scientific theory of evolution by natural selection
6765-495: Is the more common means of reproduction among eukaryotes and multicellular organisms. The Red Queen hypothesis has been used to explain the significance of sexual reproduction as a means to enable continual evolution and adaptation in response to coevolution with other species in an ever-changing environment. Another hypothesis is that sexual reproduction is primarily an adaptation for promoting accurate recombinational repair of damage in germline DNA, and that increased diversity
6888-439: Is the process by which traits that enhance survival and reproduction become more common in successive generations of a population. It embodies three principles: More offspring are produced than can possibly survive, and these conditions produce competition between organisms for survival and reproduction. Consequently, organisms with traits that give them an advantage over their competitors are more likely to pass on their traits to
7011-451: The Cambrian period. At the most basic level, predators kill and eat other organisms. However, the concept of predation is broad, defined differently in different contexts, and includes a wide variety of feeding methods; moreover, some relationships that result in the prey's death are not necessarily called predation. A parasitoid , such as an ichneumon wasp , lays its eggs in or on its host;
7134-465: The angel shark , the northern pike and the eastern frogfish . Among the many invertebrate ambush predators are trapdoor spiders and Australian Crab spiders on land and mantis shrimps in the sea. Ambush predators often construct a burrow in which to hide, improving concealment at the cost of reducing their field of vision. Some ambush predators also use lures to attract prey within striking range. The capturing movement has to be rapid to trap
7257-455: The common garter snake has developed a resistance to the toxin in the skin of the rough-skinned newt . Predators affect their ecosystems not only directly by eating their own prey, but by indirect means such as reducing predation by other species, or altering the foraging behaviour of a herbivore, as with the biodiversity effect of wolves on riverside vegetation or sea otters on kelp forests. This may explain population dynamics effects such as
7380-475: The grouper and coral trout spot prey that is inaccessible to them, they signal to giant moray eels , Napoleon wrasses or octopuses . These predators are able to access small crevices and flush out the prey. Killer whales have been known to help whalers hunt baleen whales . Social hunting allows predators to tackle a wider range of prey, but at the risk of competition for the captured food. Solitary predators have more chance of eating what they catch, at
7503-461: The marginal value theorem . Search patterns often appear random. One such is the Lévy walk , that tends to involve clusters of short steps with occasional long steps. It is a good fit to the behaviour of a wide variety of organisms including bacteria, honeybees, sharks and human hunter-gatherers. Having found prey, a predator must decide whether to pursue it or keep searching. The decision depends on
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#17330846326577626-425: The neutral theory of molecular evolution most evolutionary changes are the result of the fixation of neutral mutations by genetic drift. In this model, most genetic changes in a population are thus the result of constant mutation pressure and genetic drift. This form of the neutral theory has been debated since it does not seem to fit some genetic variation seen in nature. A better-supported version of this model
7749-454: The product of a gene , or prevent the gene from functioning, or have no effect. About half of the mutations in the coding regions of protein-coding genes are deleterious — the other half are neutral. A small percentage of the total mutations in this region confer a fitness benefit. Some of the mutations in other parts of the genome are deleterious but the vast majority are neutral. A few are beneficial. Mutations can involve large sections of
7872-519: The ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms' evolvability). Adaptation occurs through the gradual modification of existing structures. Consequently, structures with similar internal organisation may have different functions in related organisms. This is the result of a single ancestral structure being adapted to function in different ways. The bones within bat wings, for example, are very similar to those in mice feet and primate hands, due to
7995-709: The ability of predatory bacteria to digest the complex peptidoglycan polymer from the cell walls of the bacteria that they prey upon. Carnivorous vertebrates of all five major classes (fishes, amphibians, reptiles, birds, and mammals) have lower relative rates of sugar to amino acid transport than either herbivores or omnivores, presumably because they acquire plenty of amino acids from the animal proteins in their diet. To counter predation, prey have evolved defences for use at each stage of an attack. They can try to avoid detection, such as by using camouflage and mimicry . They can detect predators and warn others of their presence. If detected, they can try to avoid being
8118-728: The ability to crush or open the armoured shells of molluscs. Many predators are powerfully built and can catch and kill animals larger than themselves; this applies as much to small predators such as ants and shrews as to big and visibly muscular carnivores like the cougar and lion . Predators are often highly specialized in their diet and hunting behaviour; for example, the Eurasian lynx only hunts small ungulates . Others such as leopards are more opportunistic generalists, preying on at least 100 species. The specialists may be highly adapted to capturing their preferred prey, whereas generalists may be better able to switch to other prey when
8241-402: The adaptation of horses' teeth to the grinding of grass. By using the term adaptation for the evolutionary process and adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished. Adaptations are produced by natural selection. The following definitions are due to Theodosius Dobzhansky: Adaptation may cause either the gain of a new feature, or
8364-507: The alleles are subject to sampling error . This drift halts when an allele eventually becomes fixed, either by disappearing from the population or by replacing the other alleles entirely. Genetic drift may therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective forces, genetic drift can cause two separate populations that begin with the same genetic structure to drift apart into two divergent populations with different sets of alleles. According to
8487-584: The assault. When animals eat seeds ( seed predation or granivory ) or eggs ( egg predation ), they are consuming entire living organisms, which by definition makes them predators. Scavengers , organisms that only eat organisms found already dead, are not predators, but many predators such as the jackal and the hyena scavenge when the opportunity arises. Among invertebrates, social wasps such as yellowjackets are both hunters and scavengers of other insects. While examples of predators among mammals and birds are well known, predators can be found in
8610-684: The back (dorsal) and belly (pectoral) which lock in the erect position; as the catfish thrashes about when captured, these could pierce the predator's mouth, possibly fatally. Some fish-eating birds like the osprey avoid the danger of spines by tearing up their prey before eating it. In social predation, a group of predators cooperates to kill prey. This makes it possible to kill creatures larger than those they could overpower singly; for example, hyenas , and wolves collaborate to catch and kill herbivores as large as buffalo, and lions even hunt elephants. It can also make prey more readily available through strategies like flushing of prey and herding it into
8733-427: The beewolf's body are presented. The beewolf carries its prey back to a tunnel, but usually only stores it temporarily, until it is later used to provision a cell burrow, where an egg is laid. The tunnel of Philanthus triangulum can be as much as 1 m long. The first part of the tunnel slopes downward at an angle of 30°, after which it levels out. Up to 34 lateral tunnels, each ending in a brood chamber, branch off from
8856-441: The biosphere. For example, the overwhelming majority of species are microscopic prokaryotes , which form about half the world's biomass despite their small size and constitute the vast majority of Earth's biodiversity. Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day, with complex life only appearing more diverse because it
8979-488: The change over time in this genetic variation. The frequency of one particular allele will become more or less prevalent relative to other forms of that gene. Variation disappears when a new allele reaches the point of fixation —when it either disappears from the population or replaces the ancestral allele entirely. Mutations are changes in the DNA sequence of a cell's genome and are the ultimate source of genetic variation in all organisms. When mutations occur, they may alter
9102-416: The costs and benefits involved. A bird foraging for insects spends a lot of time searching but capturing and eating them is quick and easy, so the efficient strategy for the bird is to eat every palatable insect it finds. By contrast, a predator such as a lion or falcon finds its prey easily but capturing it requires a lot of effort. In that case, the predator is more selective. One of the factors to consider
9225-655: The cycles observed in lynx and snowshoe hares. One way of classifying predators is by trophic level . Carnivores that feed on herbivores are secondary consumers; their predators are tertiary consumers, and so forth. At the top of this food chain are apex predators such as lions . Many predators however eat from multiple levels of the food chain; a carnivore may eat both secondary and tertiary consumers. This means that many predators must contend with intraguild predation , where other predators kill and eat them. For example, coyotes compete with and sometimes kill gray foxes and bobcats . Evolution Evolution
9348-403: The descent of all these structures from a common mammalian ancestor. However, since all living organisms are related to some extent, even organs that appear to have little or no structural similarity, such as arthropod , squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes that control their assembly and function; this
9471-400: The development of thinking about the evolution of genome composition, including isochores. Different insertion vs. deletion biases in different taxa can lead to the evolution of different genome sizes. The hypothesis of Lynch regarding genome size relies on mutational biases toward increase or decrease in genome size. However, mutational hypotheses for the evolution of composition suffered
9594-490: The direct control of genes include the inheritance of cultural traits and symbiogenesis . From a neo-Darwinian perspective, evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms, for example, the allele for black colour in a population of moths becoming more common. Mechanisms that can lead to changes in allele frequencies include natural selection, genetic drift, and mutation bias. Evolution by natural selection
9717-993: The eggs hatch into larvae, which eat the host, and it inevitably dies. Zoologists generally call this a form of parasitism , though conventionally parasites are thought not to kill their hosts. A predator can be defined to differ from a parasitoid in that it has many prey, captured over its lifetime, where a parasitoid's larva has just one, or at least has its food supply provisioned for it on just one occasion. There are other difficult and borderline cases. Micropredators are small animals that, like predators, feed entirely on other organisms; they include fleas and mosquitoes that consume blood from living animals, and aphids that consume sap from living plants. However, since they typically do not kill their hosts, they are now often thought of as parasites. Animals that graze on phytoplankton or mats of microbes are predators, as they consume and kill their food organisms, while herbivores that browse leaves are not, as their food plants usually survive
9840-404: The evolution of mimicry. Avoidance is not necessarily an evolutionary response as it is generally learned from bad experiences with prey. However, when the prey is capable of killing the predator (as can a coral snake with its venom), there is no opportunity for learning and avoidance must be inherited. Predators can also respond to dangerous prey with counter-adaptations. In western North America,
9963-489: The evolutionary history of life on Earth. Morphological and biochemical traits tend to be more similar among species that share a more recent common ancestor , which historically was used to reconstruct phylogenetic trees , although direct comparison of genetic sequences is a more common method today. Evolutionary biologists have continued to study various aspects of evolution by forming and testing hypotheses as well as constructing theories based on evidence from
10086-414: The field or laboratory and on data generated by the methods of mathematical and theoretical biology . Their discoveries have influenced not just the development of biology but also other fields including agriculture, medicine, and computer science . Evolution in organisms occurs through changes in heritable characteristics—the inherited characteristics of an organism. In humans, for example, eye colour
10209-427: The fitness of an allele is not a fixed characteristic; if the environment changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become harmful. However, even if the direction of selection does reverse in this way, traits that were lost in the past may not re-evolve in an identical form. However, a re-activation of dormant genes, as long as they have not been eliminated from
10332-472: The genome and were only suppressed perhaps for hundreds of generations, can lead to the re-occurrence of traits thought to be lost like hindlegs in dolphins, teeth in chickens, wings in wingless stick insects, tails and additional nipples in humans etc. "Throwbacks" such as these are known as atavisms . Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorised into three different types. The first
10455-479: The genome of a species is very similar among all individuals of that species. However, discoveries in the field of evolutionary developmental biology have demonstrated that even relatively small differences in genotype can lead to dramatic differences in phenotype both within and between species. An individual organism's phenotype results from both its genotype and the influence of the environment it has lived in. The modern evolutionary synthesis defines evolution as
10578-640: The idea of developmental bias . Haldane and Fisher argued that, because mutation is a weak pressure easily overcome by selection, tendencies of mutation would be ineffectual except under conditions of neutral evolution or extraordinarily high mutation rates. This opposing-pressures argument was long used to dismiss the possibility of internal tendencies in evolution, until the molecular era prompted renewed interest in neutral evolution. Noboru Sueoka and Ernst Freese proposed that systematic biases in mutation might be responsible for systematic differences in genomic GC composition between species. The identification of
10701-432: The inseminated females to hunt for other invertebrates (in this case bees), on which she lays her eggs , supplying the larvae with prey when they emerge. The adults consume nectar from flowers. The prevalent European species, P. triangulum , specializes in preying upon honey bees, thus making it a minor pest for beekeepers . Other Philanthus may specialize in other bee species or they may be generalists which prey upon
10824-450: The interaction of its genotype with the environment while others are neutral. Some observable characteristics are not inherited. For example, suntanned skin comes from the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's children. The phenotype is the ability of the skin to tan when exposed to sunlight. However, some people tan more easily than others, due to differences in genotypic variation;
10947-431: The larvae of coccinellid beetles (ladybirds) , alternate between actively searching and scanning the environment. Prey distributions are often clumped, and predators respond by looking for patches where prey is dense and then searching within patches. Where food is found in patches, such as rare shoals of fish in a nearly empty ocean, the search stage requires the predator to travel for a substantial time, and to expend
11070-512: The level of the individual organism are genes called transposons , which can replicate and spread throughout a genome. Selection at a level above the individual, such as group selection , may allow the evolution of cooperation. Genetic drift is the random fluctuation of allele frequencies within a population from one generation to the next. When selective forces are absent or relatively weak, allele frequencies are equally likely to drift upward or downward in each successive generation because
11193-429: The light signals of other species, thereby attracting male fireflies, which they capture and eat. Flower mantises are ambush predators; camouflaged as flowers, such as orchids , they attract prey and seize it when it is close enough. Frogfishes are extremely well camouflaged, and actively lure their prey to approach using an esca , a bait on the end of a rod-like appendage on the head, which they wave gently to mimic
11316-452: The longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed. These outcomes of evolution are distinguished based on time scale as macroevolution versus microevolution. Macroevolution refers to evolution that occurs at or above the level of species, in particular speciation and extinction, whereas microevolution refers to smaller evolutionary changes within
11439-405: The loss of an ancestral feature. An example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters that pump the drug out of the cell. Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as
11562-413: The main tunnel. Each brood chamber is stocked with one to six honeybees. Predator Predation is a biological interaction where one organism, the predator , kills and eats another organism, its prey . It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the host ) and parasitoidism (which always does, eventually). It
11685-458: The mechanics in developmental plasticity and canalisation . Heritability may also occur at even larger scales. For example, ecological inheritance through the process of niche construction is defined by the regular and repeated activities of organisms in their environment. This generates a legacy of effects that modify and feed back into the selection regime of subsequent generations. Other examples of heritability in evolution that are not under
11808-408: The movement of pollen between heavy-metal-tolerant and heavy-metal-sensitive populations of grasses. Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer . Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria. In medicine, this contributes to
11931-427: The next generation than those with traits that do not confer an advantage. This teleonomy is the quality whereby the process of natural selection creates and preserves traits that are seemingly fitted for the functional roles they perform. Consequences of selection include nonrandom mating and genetic hitchhiking . The central concept of natural selection is the evolutionary fitness of an organism. Fitness
12054-499: The offspring of sexual organisms contain random mixtures of their parents' chromosomes that are produced through independent assortment. In a related process called homologous recombination , sexual organisms exchange DNA between two matching chromosomes. Recombination and reassortment do not alter allele frequencies, but instead change which alleles are associated with each other, producing offspring with new combinations of alleles. Sex usually increases genetic variation and may increase
12177-456: The other alleles of that gene, then with each generation this allele has a higher probability of becoming common within the population. These traits are said to be "selected for ." Examples of traits that can increase fitness are enhanced survival and increased fecundity . Conversely, the lower fitness caused by having a less beneficial or deleterious allele results in this allele likely becoming rarer—they are "selected against ." Importantly,
12300-434: The other hand, the fitness cost of a given lost dinner is unpredictable, as the predator may quickly find better prey. In addition, most predators are generalists, which reduces the impact of a given prey adaption on a predator. Since specialization is caused by predator-prey coevolution, the rarity of specialists may imply that predator-prey arms races are rare. It is difficult to determine whether given adaptations are truly
12423-578: The overall process, like a step in an assembly line. One example of mutation is wild boar piglets. They are camouflage coloured and show a characteristic pattern of dark and light longitudinal stripes. However, mutations in the melanocortin 1 receptor ( MC1R ) disrupt the pattern. The majority of pig breeds carry MC1R mutations disrupting wild-type colour and different mutations causing dominant black colouring. In asexual organisms, genes are inherited together, or linked , as they cannot mix with genes of other organisms during reproduction. In contrast,
12546-527: The presence of hip bones in whales and snakes, and sexual traits in organisms that reproduce via asexual reproduction. Examples of vestigial structures in humans include wisdom teeth , the coccyx , the vermiform appendix , and other behavioural vestiges such as goose bumps and primitive reflexes . However, many traits that appear to be simple adaptations are in fact exaptations : structures originally adapted for one function, but which coincidentally became somewhat useful for some other function in
12669-538: The prey an opportunity to escape. Some frogs wait until snakes have begun their strike before jumping, reducing the time available to the snake to recalibrate its attack, and maximising the angular adjustment that the snake would need to make to intercept the frog in real time. Ballistic predators include insects such as dragonflies, and vertebrates such as archerfish (attacking with a jet of water), chameleons (attacking with their tongues), and some colubrid snakes . In pursuit predation, predators chase fleeing prey. If
12792-549: The prey are dense and mobile, and the predator has low energy requirements. Wide foraging expends more energy, and is used when prey is sedentary or sparsely distributed. There is a continuum of search modes with intervals between periods of movement ranging from seconds to months. Sharks, sunfish , Insectivorous birds and shrews are almost always moving while web-building spiders, aquatic invertebrates, praying mantises and kestrels rarely move. In between, plovers and other shorebirds , freshwater fish including crappies , and
12915-413: The prey flees in a straight line, capture depends only on the predator's being faster than the prey. If the prey manoeuvres by turning as it flees, the predator must react in real time to calculate and follow a new intercept path, such as by parallel navigation , as it closes on the prey. Many pursuit predators use camouflage to approach the prey as close as possible unobserved ( stalking ) before starting
13038-428: The prey's body. However, the "life-dinner" principle of Dawkins and Krebs predicts that this arms race is asymmetric: if a predator fails to catch its prey, it loses its dinner, while if it succeeds, the prey loses its life. The metaphor of an arms race implies ever-escalating advances in attack and defence. However, these adaptations come with a cost; for instance, longer legs have an increased risk of breaking, while
13161-433: The prey, given that the attack is not modifiable once launched. Ballistic interception is the strategy where a predator observes the movement of a prey, predicts its motion, works out an interception path, and then attacks the prey on that path. This differs from ambush predation in that the predator adjusts its attack according to how the prey is moving. Ballistic interception involves a brief period for planning, giving
13284-497: The prey. If the attack is successful, the predator kills the prey, removes any inedible parts like the shell or spines, and eats it. Predators are adapted and often highly specialized for hunting, with acute senses such as vision , hearing , or smell . Many predatory animals , both vertebrate and invertebrate , have sharp claws or jaws to grip, kill, and cut up their prey. Other adaptations include stealth and aggressive mimicry that improve hunting efficiency. Predation has
13407-504: The price of increased expenditure of energy to catch it, and increased risk that the prey will escape. Ambush predators are often solitary to reduce the risk of becoming prey themselves. Of 245 terrestrial members of the Carnivora (the group that includes the cats, dogs, and bears), 177 are solitary; and 35 of the 37 wild cats are solitary, including the cougar and cheetah. However, the solitary cougar does allow other cougars to share in
13530-464: The process. One example is the African lizard Holaspis guentheri , which developed an extremely flat head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation. Within cells, molecular machines such as the bacterial flagella and protein sorting machinery evolved by
13653-414: The pursuit. Pursuit predators include terrestrial mammals such as humans, African wild dogs, spotted hyenas and wolves; marine predators such as dolphins, orcas and many predatory fishes, such as tuna; predatory birds (raptors) such as falcons; and insects such as dragonflies . An extreme form of pursuit is endurance or persistence hunting , in which the predator tires out the prey by following it over
13776-494: The rate of evolution. The two-fold cost of sex was first described by John Maynard Smith . The first cost is that in sexually dimorphic species only one of the two sexes can bear young. This cost does not apply to hermaphroditic species, like most plants and many invertebrates . The second cost is that any individual who reproduces sexually can only pass on 50% of its genes to any individual offspring, with even less passed on as each new generation passes. Yet sexual reproduction
13899-408: The relative importance of selection and neutral processes, including drift. The comparative importance of adaptive and non-adaptive forces in driving evolutionary change is an area of current research . Mutation bias is usually conceived as a difference in expected rates for two different kinds of mutation, e.g., transition-transversion bias, GC-AT bias, deletion-insertion bias. This is related to
14022-470: The result of coevolution, where a prey adaptation gives rise to a predator adaptation that is countered by further adaptation in the prey. An alternative explanation is escalation , where predators are adapting to competitors, their own predators or dangerous prey. Apparent adaptations to predation may also have arisen for other reasons and then been co-opted for attack or defence. In some of the insects preyed on by bats, hearing evolved before bats appeared and
14145-417: The same strand of DNA to become separated. However, the rate of recombination is low (approximately two events per chromosome per generation). As a result, genes close together on a chromosome may not always be shuffled away from each other and genes that are close together tend to be inherited together, a phenomenon known as linkage . This tendency is measured by finding how often two alleles occur together on
14268-552: The sequence of nucleotides in the DNA. These phenomena are classed as epigenetic inheritance systems. DNA methylation marking chromatin , self-sustaining metabolic loops, gene silencing by RNA interference and the three-dimensional conformation of proteins (such as prions ) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in genetic networks and communication among cells can lead to heritable variations that may underlay some of
14391-417: The specialized tongue of the chameleon, with its ability to act like a projectile, is useless for lapping water, so the chameleon must drink dew off vegetation. The "life-dinner" principle has been criticized on multiple grounds. The extent of the asymmetry in natural selection depends in part on the heritability of the adaptive traits. Also, if a predator loses enough dinners, it too will lose its life. On
14514-406: The spread of antibiotic resistance , as when one bacteria acquires resistance genes it can rapidly transfer them to other species. Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean weevil Callosobruchus chinensis has occurred. An example of larger-scale transfers are the eukaryotic bdelloid rotifers , which have received
14637-640: The survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these hard-to-fake , sexually selected traits. Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical adaptations that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by cooperating with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis . In
14760-642: The system...." Each population within an ecosystem occupies a distinct niche , or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the food chain and its geographic range. This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection. Natural selection can act at different levels of organisation , such as genes, cells, individual organisms, groups of organisms and species. Selection can act at multiple levels simultaneously. An example of selection occurring below
14883-684: The target of an attack, for example, by signalling that they are toxic or unpalatable , by signalling that a chase would be unprofitable, or by forming groups. If they become a target, they can try to fend off the attack with defences such as armour, quills , unpalatability, or mobbing; and they can often escape an attack in progress by startling the predator, playing dead , shedding body parts such as tails, or simply fleeing. Predators and prey are natural enemies, and many of their adaptations seem designed to counter each other. For example, bats have sophisticated echolocation systems to detect insects and other prey, and insects have developed
15006-887: Was conceived independently by two British naturalists, Charles Darwin and Alfred Russel Wallace , in the mid-19th century as an explanation for why organisms are adapted to their physical and biological environments. The theory was first set out in detail in Darwin's book On the Origin of Species . Evolution by natural selection is established by observable facts about living organisms: (1) more offspring are often produced than can possibly survive; (2) traits vary among individuals with respect to their morphology , physiology , and behaviour; (3) different traits confer different rates of survival and reproduction (differential fitness ); and (4) traits can be passed from generation to generation ( heritability of fitness). In successive generations, members of
15129-401: Was used to hear signals used for territorial defence and mating. Their hearing evolved in response to bat predation, but the only clear example of reciprocal adaptation in bats is stealth echolocation. A more symmetric arms race may occur when the prey are dangerous, having spines, quills, toxins or venom that can harm the predator. The predator can respond with avoidance, which in turn drives
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