Inevolutionary biology,mimicry in plants is where a plant evolves to resemble another organism physically or chemically. Mimicry in plants has been studied far less thanmimicry in animals. It may provideprotection against herbivory, or may deceptively encourage mutualists, likepollinators, to provide a service without offering a reward in return.[2]
Types of plant mimicry includeBakerian, where female flowers imitate males of the same species;Dodsonian, where a plant mimics a rewarding flower, luring pollinators by mimicking another species of flower, or fruit where feeders of the other species are attracted to a fake fruit to distribute seeds;Gilbertian, where a plant has structures like butterfly eggs, dissuading egg-laying;Vavilovian, where aweed isunintentionally selected to resemble acrop plant;Pouyannian, in which a flower imitates a female mate, deceiving a male pollinating insect intopseudocopulation;Batesian, where a harmless species deter predators by mimicking the characteristics of a harmful species; and leaf mimicry, where a plant resembles a nearby plant to evade the attention of herbivores.
Mimicry is anadaptation by a species, called the mimic, making it resemble something else, called the model, with the effect of deceiving another species, the dupe. The three are not always all distinct, as mimicry can for example be within a species.[3] The adaptation is to theevolutionary advantage of the mimic. As such it can be any mechanism that may evolve bynatural selection; no conscious intention is involved. There is no essential difference between the evolution of mimicry in plants and in other organisms such asin animals, though mimicry in animals is better known. Plant mimicry can broadly be divided into reproductive mimicry that deceivespollinators into providing their service without rewarding them in return, and protective mimicry that deceives herbivores into not eating the plant.[2]
Mimicry mechanisms, including theBatesian[4] andMüllerian[5] mimicry, were described in animals in the 19th century.[3]Charles Darwin's 1862 bookFertilisation of Orchids laid the foundations for research into plantreproductive strategies co-evolved with insects. In the book, Darwin wonders why some orchids apparently mimic bees, and how orchids without nectar succeed in attracting pollinators.[6] Several mimicry mechanisms in plants were described in the 20th century, starting withPouyannian (1916),[7] Vavilovian (1951),[8] andGilbertian (1975).[9] Study of mimicry in plants broadened in the 21st century, with the discovery of Batesian mimicry in plants in 2003,[10] and crypsis or leaf mimicry in 2014.[1] Until then, mimicry in plants was studied infrequently, both because it is uncommon and because botanists considered mainly physical factors in plant ecology. It may be that since plants often grow in clusters, foraging herbivores are hard to deceive once they have started to feed in an area.[11]
Carrion flowers, including the enormousAmorphophallus titanum,[11] mimic the scent and appearance of rotting flesh to attract necrophagous (carrion-feeding) insects like flesh flies (Sarcophagidae), blowflies (Calliphoridae), house flies (Muscidae) and some beetles (e.g.,Dermestidae andSilphidae) which search for dead animals to use as brood sites.[12][13] The decaying smell of the flower comes fromoligosulfides, compounds found in decayed proteins with the sulfur-containing amino acidsmethionine andcysteine. While carrion flowers do produce a small amount ofnectar, this does not necessarily make their relationship to necrophagous insectsmutualistic. Insects lay eggs on the carrion flowers, meaning they mistake the flowers foroviposition sites. The nectar acts as a lure to bring the insects closer to the reproductive parts of the flower.[12]
Bakerian mimicry, named after English naturalistHerbert Baker,[14] is a form ofautomimicry orintraspecific mimicry that occurs within a single species. In plants, the femaleflowers mimic male flowers of their own species, cheating pollinators out of a reward. This reproductive mimicry may not be readily apparent as members of the same species may still exhibit some degree ofsexual dimorphism, i.e. thephenotypic difference between males and females of the same species. It is common in many species ofCaricaceae, afamily offlowering plants in the orderBrassicales, found primarily intropical regions ofCentral andSouth America, andAfrica.[15][16]
Dodsonian mimicry, named after the AmericanbotanistCalaway H. Dodson, is a form of reproductive floral mimicry where the model belongs to a different species than the mimic. By providing similar sensory signals to the model's flowers, the mimic lures the model's pollinators. Like Bakerian mimics, nonectar is provided.[17]
Epidendrum ibaguense, a species ofepiphytic orchid of the genusEpidendrum that occurs in the northern part of South America, resembles flowers ofLantana camara andAsclepias curassavica.Epidendrum ibaguense is pollinated bymonarch butterfly (Danaus plexippus) and perhapshummingbirds.[18] Similar cases are seen in some other species of the same family. The mimetic species may still have pollinators of its own; for example, aLamellicorn beetle helps to pollinateOphrys species mainly pollinated bybees.[19]
InPouyannian mimicry,[20][7] named after the French lawyer and amateur botanist Maurice-Alexandre Pouyanne,[21] flowers mimic a malepollinator's potential female mate, visually or with other stimuli.[22] Many orchids, including theOphrys bee orchids, deceive male insects intopseudocopulation in this way, using them to transfer pollen.[23] For instance, the orchidEpipactis helleborine is physiologically andmorphologically adapted to attractsocial wasps as their primary pollinators. Social wasps feed their larvae on insects such as caterpillars. To locate that prey, they use a combination of visual andolfactory cues. The flowers ofE. helleborine andE. purpurata emit green-leafvolatiles attractive to foragers of the social waspsVespula germanica andV. vulgaris.E. helleborine emits several green-leaf volatiles that induce a response in wasp antennae. These same volatiles are also produced by cabbage leaves infested with caterpillars (Pieris brassicae), which are common prey items for wasps. Despite a large nectar reward, the species is almost entirely overlooked by other pollinators.[24]
In Batesian mimicry, named after the English naturalistHenry Walter Bates, a harmless species has evolved to imitate the warning signals of a harmful species directed at apredator. Batesian mimicry ofthorns[10] and possibly of spider webs[25] has been observed in plants.
Thorn mimicry of two types has been observed in plants. The first, a special case of intra-organismic Batesian mimicry characteristic ofAloe species (Liliaceae),Washingtonia filifera (Arecaceae), and dozens of species ofAgave, includingA. applanta,A. salmiana, andA. obscura. These plants develop thornlike imprints or colorations on the face of their leaves due to the teeth along the margins of that leaf (or another leaf) pressing sustained indentations into the flesh of the non-spiny parts. The second type of thorn mimicry, a more classic case of Batesian mimicry, involves the pointed, colorful organs likebuds, leaves and fruit of mimetic plant species that mimic warning-colouredaposematic colorful thorns not found anywhere else in the organism.[10]
Several plants from different parts of the world may be mimics of spider webs. Dense, whitetrichomes are produced on newly extended stems and leaves that deterherbivory due to predatory habit or toxicity. This may be a case of visual mimicry or perceptual exploitation. Case examples include the new buds ofOnopordum from Israel,Carthamus species from Greece, flower heads ofArctium tomentosum from Estonia, a fledgling leaf ofTussilago farfara from Estonia, and newfronds ofOsmunda japonica from Japan.[25]
Another plant leaf pattern has been suggested to be mimetic: irregular white blotches on leaves of plants such asPulmonaria officinalis perhaps protect against large herbivores through their resemblance to bird droppings, which could be avoided as possible sources of disease.[2]
Gilbertian[26] or colonisation mimicry[2] is bipolar, involving only two species. The potential host (or prey) drives away its parasite (or predator) by mimicking it, the reverse of host–parasite aggressive mimicry. Georges Pasteur named it after the AmericanecologistLawrence E. Gilbert, who described it in 1975.[26][9] The classical instance of Gilbertian mimicry is in the plant genusPassiflora, which is grazed by themicropredator larvae of someHeliconius butterflies. The host plants have evolvedstipules, small outgrowths at the base of each leaf, that mimic matureHeliconius eggs near the point of hatching. The butterflies avoid laying eggs near existing ones, reducingintraspecific competition between caterpillars, which are alsocannibalistic, so those that lay on vacant leaves provide their offspring with a greater chance of survival. The stipules thus appear to have evolved as Gilbertian mimics of butterfly eggs, under selection pressure from these caterpillars.[26][27]
In ecology,crypsis is an organism's ability to avoid detection by other organisms. In cryptic mimicry, a prey organism deceives a potential predator by providing false, usually visual, signals or a lack of signals, with the effect ofcamouflaging itself.Boquila trifoliata, a South American member of the familyLardizabalaceae, is a climbing vine with a highly variable appearance (phenotype). It is capable of mimicking the leaf features of plant species that it clings to, such as their coloration, size, and shape. By camouflaging its leaves,Boquila reduces damage from herbivorous animals.[1] It has been speculated that such plants may make use of "some kind of vision" using ocelli, or "delicate chemical sensing", to account for the mimic's ability to cope with such a large number of variables in its model's appearance,[29] including the ability to mimic the foliage of an artificial host plant made of plastic.[30] Another plant that could well be a cryptic mimic of its host is the parasitic Australian mistletoe,Amyema cambagei, which has an "uncanny resemblance" to the foliage ofCasuarina trees.[2]
Some hundreds of species in theMesembryanthemaceae (ice plants) of Southern Africa arecamouflaged as small stones, especiallyLithops, which are known as pebble plants or living stones. By appearing non-living, they are less likely to be eaten by herbivores, and in dusty dry conditions among stones are extremely difficult to detect.[11]
Vavilovian mimicry (also known as crop mimicry or weed mimicry[31]) is named after the Russian plant geneticist who identified thecentres of origin of cultivated plants,Nikolai Vavilov.[8] It is a form of mimicry in plants where aweed comes to share one or more characteristics with adomesticated plant through generations ofunintentional selection. Selection against the weed may occur by killing a young or adult weed, separating itsseeds from those of the crop (winnowing), or both. This has been done manually sinceNeolithic times, and in more recent years byagricultural machinery.[32]
Darwin's open questions about orchid fertilisation stimulated popularizers such asGrant Allen to write on the topic, and led thescience fiction writerH. G. Wells to tell a tale of killer orchids, his 1896 "The Flowering of the Strange Orchid". The vampire-like plant attacks people using "tentacle-like aerial rootlets". Other authors followed with similar tales of their own.[6][33]