Pouyannian Mimicry
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Pouyannian mimicry is a form of
mimicry in plants In evolutionary biology, mimicry in plants is where a plant evolves to resemble another organism physically or chemically. Mimicry in plants has been studied far less than mimicry in animals. It may provide protection against herbivory, or may ...
that deceives an insect into attempting to copulate with a flower. The flower mimics a potential female mate of a male insect, which then serves the plant as a
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
. The mechanism is named after the French lawyer and amateur botanist Maurice-Alexandre Pouyanne. The resemblance that he noted is visual, but the key stimuli that deceive the pollinator are often chemical and tactile. In
orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan plants that are found in almost every habitat on Eart ...
s, the resemblance is to a species of bee; Pouyanne observed the bee ''
Dasyscolia ciliata ''Dasyscolia ciliata'' is a species of scoliid wasp found throughout the Mediterranean. It is the only species in the genus ''Dasyscolia''. It is the only known pollinator of the European ''Ophrys speculum''. The male wasp is tricked into pollina ...
'' pollinating the orchid ''
Ophrys speculum ''Ophrys speculum'', the mirror orchid, is a species of ''Ophrys'' distributed throughout the Mediterranean region, Mediterranean that is pollinated exclusively by a single species of scoliid wasp. Description A terrestrial orchid up to 25  ...
''. The flower uses morphology, coloration, and scent to deceive the pollinator. The chemicals secreted from the flower's osmophore glands are indistinguishable from the insect's pheromones. The pollinator is not rewarded with nectar, and may waste significant amounts of sperm while trying to mate with the flower.


Definition

The form of
mimicry in plants In evolutionary biology, mimicry in plants is where a plant evolves to resemble another organism physically or chemically. Mimicry in plants has been studied far less than mimicry in animals. It may provide protection against herbivory, or may ...
that deceives an insect into
pseudocopulation Pseudocopulation is a behavior similar to Copulation (zoology), copulation that serves a reproductive function for one or both participants but does not involve actual sexual union between the individuals. It is most generally applied to a pollin ...
is called Pouyannian mimicry after the French lawyer and amateur botanist Maurice-Alexandre Pouyanne.
Mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
typically involves three species, namely a mimic, a model, and a dupe, as seen for example in
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
. Floral mimicry involves the imitation of other plants or animals, including of coloration, morphology, egg deposition sites, provoking scents, and mating signals. In the case of Pouyannian mimicry, the model and the dupe are the female and male of the same species, so the mimicry is bipolar, involving only two species, an insect and a flower.


History

In 1916, Pouyanne, with Henry Correvon, described his observations in Algeria:


In orchids

Several
orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan plants that are found in almost every habitat on Eart ...
s (''Orchidaceae'') make use of floral mimicry. Using sex-based deception, these species imitate female mating signals of certain pollinator species. This results in attempted copulation by males of the pollinator species, facilitating pollen transfer. Bee orchids (''
Ophrys apifera ''Ophrys apifera'', known in Europe as the bee orchid, is a perennial plant, perennial herbaceous plant of the genus ''Ophrys'', in the family of Orchidaceae. It serves as an example of Sexual mimicry#Interspecific deceptive mimicry, sexually d ...
'') and fly orchids (''
Ophrys insectifera ''Ophrys insectifera'', the fly orchid, is a species of orchid and the type species of the genus ''Ophrys''. It is remarkable as an example of the use of sexually deceptive pollination and floral mimicry, as well as a highly selective and highly ...
''), specifically, utilize flower morphology, coloration, and scent to deceive their respective pollinators. These orchids have evolved traits matching the preferences of specific pollinator niches, leading to adaptive
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. The mimicry involves secreting chemicals from glands (osmophores) in the
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s,
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s, or labellum, that are indistinguishable from the insect's natural
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s. The flower attaches a
pollinium A pollinium (: pollinia) is a coherent mass of pollen grains in a plant that are the product of only one anther, but are transferred, during pollination, as a single unit. This is regularly seen in plants such as orchid Orchids are plants tha ...
to the pollinator's body; the insect transfers the pollinium to the stigma of another flower when it makes its next copulation attempt. Pollinators are often
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s, or
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
. The cost to the pollinating insects might be seen as negligible, but pollinators of the Australian orchid ''
Cryptostylis ''Cryptostylis'', commonly known as tongue orchids, is a genus of flowering plants from the orchid family. Tongue orchids are terrestrial herbs with one to a few stalked leaves at the base of the flowering stem, or leafless. One to a few dull co ...
'' can waste significant amounts of
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
by ejaculating onto the flower. Thus there could be antagonistic
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
such that pollinators become better at identifying their own species correctly, while orchids become better mimics. File:Ophrys apifera - panoramio (52).jpg, Mimic: flower of Bee Orchid, ''
Ophrys apifera ''Ophrys apifera'', known in Europe as the bee orchid, is a perennial plant, perennial herbaceous plant of the genus ''Ophrys'', in the family of Orchidaceae. It serves as an example of Sexual mimicry#Interspecific deceptive mimicry, sexually d ...
'' mimics a female bee to attract male bees as pollinators. File:GL.109 Eucera longicornis f.jpg, Model: Female longhorn bee, ''
Eucera longicornis ''Eucera longicornis'' is a species of bee in the family Apidae, subfamily Apinae, and tribe Eucerini, the long-horned bees. Description The head, thorax and abdomen of this species are black, the thorax being clad in pale brown, downy hairs on ...
''
One mechanism in pollination is to use incentives or rewards. These are beneficial offerings to a pollinator, enticing it to engage with the reward and thus transfer pollen. Flowering plants that do not produce such rewards can instead attract pollinators through mimicry — a form of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
. Such plants are called "deceptive plants" as they mimic the characteristics or rewards of other species without providing any benefit to the pollinator. Although bee and fly orchids are visual mimics of their pollinators, visual traits are not the only (nor the most important) ones mimicked to increase attraction. Floral odours have been identified as the most prominent way of attracting pollinators, because these odours imitate the
sex pheromone Sex pheromones are pheromones released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with sexual reproduction. Sex pheromones specifically focus on ind ...
s of females of the pollinator species. Male pollinators then track these scents over long distances. The proportions of such odour compounds have been found to be varied in different populations of orchids (in a variety of locations), playing a crucial role in attracting specific pollinators at the population level. The evolution of these interactions between plants and pollinators involves natural selection favoring local adaptation, leading to a more precise imitation of the scents produced by local pollinators. Chemical compounds (more specifically,
alkane In organic chemistry, an alkane, or paraffin (a historical trivial name that also has other meanings), is an acyclic saturated hydrocarbon. In other words, an alkane consists of hydrogen and carbon atoms arranged in a tree structure in whi ...
s and
alkene In organic chemistry, an alkene, or olefin, is a hydrocarbon containing a carbon–carbon double bond. The double bond may be internal or at the terminal position. Terminal alkenes are also known as Alpha-olefin, α-olefins. The Internationa ...
s), while used for sexual deception, are produced in many species of ''Ophrys'', and likely were preadapted for other functions before being co-opted for mimicry. These orchids increased ancestral levels of alkene production to mimic the female pheromones that attract male pollinators, a form of sensory exploitation called a sensory trap. Although mimetic plants typically receive fewer interactions with pollinators than truly-rewarding plants do, the evolution of sexual deception appears to be linked to benefits associated with mating behavior. Sex-based mimicry results in pollinator fidelity, the continued revisiting of flowers of the same species by a pollinator, as a result of sexual deception. In support of this, sex-based deception in an Australian orchid results in a higher proportion of pollen reaching stigmas than food-based deception. In another study, deception of male pollinators results in a long-distance dispersal of pollen.


References

{{Mimicry Mimicry Animal sexuality Pollination