Boquila

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Boquila
Boquila trifoliolata (Valdivia, Chili).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Phylum: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Ranunculales
Family: Lardizabalaceae
Genus: Boquila
Decne.
Species:
B. trifoliolata
Binomial name
Boquila trifoliolata
Synonyms [1]
  • Boquila discolor(Kunze ex Poepp. & Endl.) Decne.
  • Dolichos funarius(Molina)
  • Lardizabala funaria(Looser)
  • Lardizabala trifoliolataDC.

Boquila is a genus of flowering plants in the family Lardizabalaceae, endemic to temperate forests of central and southern Chile and Argentina. It is monotypic, being represented by the single species Boquila trifoliolata, locally known as voqui blanco or pilpil in its native range, [2] and sometimes referred as the chameleon vine since a recent report on leaf mimicry. The species was first described in 1782 by Juan Ignacio Molina, and the genus itself was established in 1839 by Joseph Decaisne. B. trifoliata forms non-parasitic vines that wind around host plants, using them for structure and protection. B. trifoliata is monoecious, and its flowers are an off white color. It bears an edible fruit and has been historically used in rope and basket making.

Contents

B. trifoliata is the only known plant species reported to engage in mimetic polymorphism, or the ability to mimic multiple host species, often simultaneously. This is a form of Batesian mimicry, when a harmless species mimics a harmful one to ward off predators. Contact between the vines and host trees was reported not to be necessary for mimicking to commence. However, after a decade of the original study describing the species mimicry capabilities in 2014, no independent research groups have verified the field observations and the mechanism by which this mimicry would occur is still unknown. Hypotheses about the mimicry mechanism include microbial mediated horizontal gene transfer, volatile organic compound sensing, and the use of eye-like structures.

Taxonomy and etymology

Boquila is a monotypic genus of flowering plants (angiosperms) in the family Lardizabalaceae with one known species, Boquila trifoliolata. [3] [4] The species was first described as Dolichosfunarius in 1782 by Juan Ignacio Molina, and in 1817, the holotype Lardizabalatrifoliolata was named by Augustin Pyramus de Candolle. [5] [6] In 1838, Stephan Endlicher, Eduard Friedrich Poeppig, and Gustav Kunze proposed the name Lardizabaladiscolor. In between 1837 to 1839, Joseph Decaisne identified Boquilatrifoliolata and Boquiladiscolor, and established the Boquila genus in 1837. [7] The name Boquiladiscolor was later declared a orthographic variant. In 1936, Gualterio Looser attempted to reclassify the species to Lardizabalafunaria based upon the observations of Carlo Giuseppe Bertero, but this classification is not considered valid. [8] [9]

Due to its mimicry capabilities, Boquila trifoliolata is sometimes referred to as the chameleon vine. [10] [11] [12]

Description

Illustration of Boquila trifoliolata by Pierre Jean Francois Turpin Boquila trifoliolata.jpg
Illustration of Boquila trifoliolata by Pierre Jean François Turpin
Interior cross-section of the Boquila trifoliolata flower Boquila trifoliolata (Parque Nacional Puyehue, Chili) 2.jpg
Interior cross-section of the Boquila trifoliolata flower

Boquila trifoliolata is a woody vine with a highly variable appearance due to its crypsis abilities. [4] The vines are evergreen or partly deciduous, meaning they largely retain their leaves over winter. [13] The vines follow a twining pattern when climbing host plants, meaning the stems bend around host plants during their ascent. [14] The branches are thin, less than 1 cm (0.39 in) in diameter, and are covered in red-brown bark. The lenticels are elliptical in shape, and the wider branches are a speckled grey color. [13] When not mimicking a host plant, B. trifoliata employs smaller 'charlatan leaves' that are short, stubby, and have three lobes (trifoliate). [4] The petioles range from 2 cm (0.79 in) to 6 cm (2.4 in) in length and the petiolules range from 0.5 cm (0.20 in) to 1.5 cm (0.59 in) in length. Leaflets are oval or elliptical and range from 2 cm (0.79 in) to 6 cm (2.4 in) in height and 1 cm (0.39 in) to 3 cm (1.2 in) in width. The base of the leaves is rounded, the margins are irregular (most often trilobate), the tips are rounded and wide-angled, the top of the leaves are dark green and hairless, the undersides are glaucous (pale-grey to blue-green), and the veins have a pinnate pattern. [15] [13]

Reproduction

In its natural habitat, flowering occurs between September and December, while fruiting occurs between January and March. This pattern is opposite when the plant is raised in the Northern Hemisphere. [13] B. trifoliata is monoecious, meaning that both male and female floral parts are present in the same plant. [13] The petals are small (1.5 cm (0.59 in) to 3 cm (1.2 in) in length) and have a green-white to yellow-white color. These flowers tend to be in 2- to 4-flower umbels with small hairs and lepidote bracts along the petals. Each flower has six sepals, and are biserate, petaloid, ovate, and the three inner sepals are larger than the outer ones. Staminate flowers (male flowers) have six stamens, petals in an opposite pattern, and anthers are oblate. Carpellate flowers (female flowers) have six conical staminodes, three carpels, an elongated stigma, and sutures running vertically up the petals. [15] [13]

The fruits are small, ranging from 0.5 cm (0.20 in) to 1 cm (0.39 in) in diameter, and white. There are typically 1-4 seeds per berry, ranging from 2.5 mm (0.098 in) to 5 mm (0.20 in). The seeds are oval, brown, and contain large amounts of endosperm. [13] Seeds are largely dispersed via animal vectors and readily germinate when planted. [15] [16]

Mimicry

Boquila trifoliolata is the only plant known to engage in mimetic polymorphism, meaning it can mimic the leaves of multiple host plants. [4] [17] Other species of vines are capable of limited crypsis for one host species, but B. trifoliata is notable since it can mimic the leaves of multiple species, with one vine capable of simultaneously mimicking multiple hosts. Mimetic polymorphism is only observed elsewhere in some species of butterflies, but that is the result of genetic divergence, unlike B. trifoliata which engages in rapid changes in leaf morphology. [4]

Once the vines approach a host tree's branches, the leaves begin to change their size, shape, color, vein patterns, spines, and orientation to match the host plant; sometimes expanding to 10x their original size. [4] B. trifoliata has been observed mimicking over 20 different species of plants. [18] These include native species such as Luma apiculata , Cissus striata , and Rhaphithamnus spinosus but also non-native species such as Ranunculus repens . [19]

Unlike most other mimicking species, close proximity is enough to induce mimicry and contact is not required. [4] In one controversial study, B. trifoliata has been noted to mimic the leaves of plastic plants. [20] If the vines approach another tree, the vine begins simultaneously mimicking that species as well. [4] Mimicry is largely confined to the leaves closest to the host, meaning that sections of the vine approximately 60 cm (24 in) away from the host retain the non-mimicking phenotype. [21] This is a form of Batesian mimicry, where the B. trifoliata is harmless but resembles a less palatable or harmful plant to ward off herbivory species and pests. [22] [23] [4]

Possible explanations

The exact mechanism by which mimicry occurs is not well understood but may involve chemical, odor, genetic, metagenomic, transcriptomic, proteomic, metabolomic, epigenetic, and/or microbial cues to identify and mimic the species it is attached to. [21] [4]

Volatile organic compounds

Plant ecologist Ernesto Gianoli proposed that the host tree may be emitting volatile organic compounds (VOCs) into the environment that B. trifoliata can detect. [4] The use of VOC-mediated plant-to-plant communication is widely employed in non-specific biological processes, including up-regulation of defense-related genes, and could explain why no contact is necessary for mimicry. Criticisms of this hypothesis are that this would mark the first time that VOCs were used to change plant morphology, and that B. trifoliata's mimicry has a level of specificity that is not normally seen with VOC-mediated responses. [21]

Horizontal gene transfer

Another hypothesis proposed by Gianoli is that B. trifoliata's mimicry is mediated by endophytic microbes that conduct horizontal gene transfer (HGT) between B. trifoliata and the host plant. This would influence the genes, transposons, and/or epigenetics of the plant's leaves, identifying the host and changing the leaf's morphology without necessitating physical contact. [19] [24] In a 2021 study, Gianoli found that the microbiomes of B. trifoliata and its host plant show significant overlap following the initiation of mimicry. Gianoli has argued this could represent a mechanism behind B. trifoliata's mimicry but still acknowledged that there are limitations to this hypothesis. While HGT commonly occurs between different species, it takes many years and manifests in discrete events. Additionally, HGT between plants is most commonly observed in cases of parasitism, which B. trifoliata does not engage in. [19]

Ocelli

In a 2021 study published in the journal Plant Signaling & Behavior, Felipe Yamashita and Jacob White claimed that B. trifoliata may employ a primitive form of vision to identify and mimic their hosts. This hypothesis is based upon 1905 and 1907 claims by Gottlieb Haberlandt and Francis Darwin, respectively, that some plants use 'ocelli' or lens-like cells to focus light onto other light sensitive cells. In this study, B. trifoliata was observed mimicking the leaf shapes of plastic plants, and researchers refined Haberlandt and Darwin's ocelli hypothesis, claiming that B. trifoliata may be using convex shaped lenses in epidermal tissue that can detect light and "see" the shapes of nearby leaves. [25] They further proposed that, B. trifoliata processes that information through an unknown means, possibly through neuron-like structures in order to initiate mimicry. [18] [24] The study also found that non-mimetic leaves have more free-end veinlets and identified the hormone auxin as a possible mediator in changes to leaf morphology. [25]

This paper received substantial media coverage, was praised by F1000's Faculty Opinions, and went viral on the social media platform TikTok following its release. Plant biologist and Plant Signaling and Behavior's editor-in-chief, František Baluška praised this hypothesis, and claimed that root skototropism and photoreceptive cells in algae were analogous mechanisms for "plant sight". However, the paper's conclusions have largely been met with skepticism by scientists. Criticisms of the paper include poor methodology, White's lack of a scientific background, and possible conflicts of interest between Baluška and Yamashita. [18] [24]

Distribution and habitat

Flowering specimen of Boquila trifoliolata at Puyehue National Park Boquila trifoliolata (Parque Nacional Puyehue, Chili) 1.jpg
Flowering specimen of Boquila trifoliolata at Puyehue National Park

The Boquila genus is endemic to the temperate rainforests, nothofagus forests, and evergreen forests of southern Argentina and Chile, ranging from Cauquenes to Chiloe. [13] [4] B. trifoliata is most commonly found between 100 metres (330 ft) to 600 metres (2,000 ft) in elevation. [13] Unlike many other species of vines, B. trifoliata is not parasitic. Instead, it only attaches to trees for protection and structure, sometimes forming thickets over 6 metres (20 ft) in height. B. trifoliata can survive temperatures as low as −8 °C (18 °F) and prefers soil rich in humus. The species is resistant to wilting, but generally prefers to grow in shaded environments. [4] [15] [13]

Human uses

The stems are used locally in basketry and in rope making. The leaf juice was historically used by local tribes to treat sore eyes and was once believed to be an aphrodisiac. The plant is also used ornamentally and the berries are edible. [13] Stems are often cut in the summer and rooted in cold frames as a means of propagation. [15]

See also

Related Research Articles

<span class="mw-page-title-main">Vine</span> Plant with a growth habit of trailing or scandent stems or runners

A vine is any plant with a growth habit of trailing or scandent stems, lianas, or runners. The word vine can also refer to such stems or runners themselves, for instance, when used in wicker work.

<span class="mw-page-title-main">Mimicry</span> Imitation of another species for selective advantage

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. Often, mimicry functions to protect a species from predators, making it an anti-predator adaptation. Mimicry evolves if a receiver perceives the similarity between a mimic and a model and as a result changes its behaviour in a way that provides a selective advantage to the mimic. The resemblances that evolve in mimicry can be visual, acoustic, chemical, tactile, or electric, or combinations of these sensory modalities. Mimicry may be to the advantage of both organisms that share a resemblance, in which case it is a form of mutualism; or mimicry can be to the detriment of one, making it parasitic or competitive. The evolutionary convergence between groups is driven by the selective action of a signal-receiver or dupe. Birds, for example, use sight to identify palatable insects and butterflies, whilst avoiding the noxious ones. Over time, palatable insects may evolve to resemble noxious ones, making them mimics and the noxious ones models. In the case of mutualism, sometimes both groups are referred to as "co-mimics". It is often thought that models must be more abundant than mimics, but this is not so. Mimicry may involve numerous species; many harmless species such as hoverflies are Batesian mimics of strongly defended species such as wasps, while many such well-defended species form Müllerian mimicry rings, all resembling each other. Mimicry between prey species and their predators often involves three or more species.

<i>Ptelea trifoliata</i> Species of tree

Ptelea trifoliata, commonly known as common hoptree, wafer ash, stinking ash, and skunk bush, is a species of flowering plant in the citrus family (Rutaceae). It is native to North America, where it is found in Canada, Mexico, and the United States. It is a deciduous shrub or tree, with alternate, trifoliate leaves.

<span class="mw-page-title-main">Batesian mimicry</span> Bluffing imitation of a strongly defended species

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 butterflies in the rainforests of Brazil.

<span class="mw-page-title-main">Viceroy (butterfly)</span> Species of butterfly

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<span class="mw-page-title-main">Müllerian mimicry</span> Mutually beneficial mimicry of strongly defended species

Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878, supporting his theory with the first mathematical model of frequency-dependent selection, one of the first such models anywhere in biology.

<span class="mw-page-title-main">Crypsis</span> Aspect of animal behaviour and morphology

In ecology, crypsis is the ability of an animal or a plant to avoid observation or detection by other animals. It may be a predation strategy or an antipredator adaptation. Methods include camouflage, nocturnality, subterranean lifestyle and mimicry. Crypsis can involve visual, olfactory or auditory concealment. When it is visual, the term cryptic coloration, effectively a synonym for animal camouflage, is sometimes used, but many different methods of camouflage are employed by animals or plants.

<span class="mw-page-title-main">Lardizabalaceae</span> Family of flowering plants

Lardizabalaceae is a family of flowering plants.

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<span class="mw-page-title-main">Aggressive mimicry</span> Deceptive mimicry of a harmless species by a predator

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<span class="mw-page-title-main">Mimicry in plants</span>

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References

  1. ↑ "The Plant List: A Working List of All Plant Species". Archived from the original on May 24, 2019. Retrieved June 19, 2014.
  2. ↑ "pilpil (Boquila trifoliolata)". iNaturalist. Retrieved March 22, 2024.
  3. ↑ taxonomy. "Taxonomy browser (Boquila trifoliolata)". www.ncbi.nlm.nih.gov. Archived from the original on December 9, 2023. Retrieved December 9, 2023.
  4. 1 2 3 4 5 6 7 8 9 10 11 12 13 Gianoli, Ernesto; Carrasco-Urra, Fernando (2014). "Leaf Mimicry in a Climbing Plant Protects against Herbivory". Current Biology. 24 (9): P984-987. Bibcode:2014CBio...24..984G. doi: 10.1016/j.cub.2014.03.010 . PMID   24768053.
  5. ↑ Molina, Giovanni Ignazio (1782). "Saggio sulla storia naturale del Chili". Biodiversity Heritage Library. Retrieved January 4, 2024.
  6. ↑ Candolle, Augustin Pyramus de; Candolle, Augustin Pyramus de (1818). Regni vegetabilis systema naturale, sive Ordines, genera et species plantarum secundum methodi naturalis normas digestarum et descriptarum. Vol. 1. Parisiis [etc.]: sumptibus sociorum Treuttel et Würtz.
  7. ↑ Compt. Rend. Hebd. Séances Acad. Sci. 3: 394 (1837)
  8. ↑ Lista Pls. Obs. Chile en 1828 por Bertero 36. 1936.
  9. ↑ Zuloaga, F. O., O. Morrone, M. J. Belgrano, C. F. S. Marticorena & E. Marchesi. (eds.) 2008. Catálogo de las plantas vasculares del Cono Sur. Monogr. Syst. Bot. Missouri Bot. Gard. 107(1–3)
  10. ↑ Yang, Ina (May 7, 2014). "'Chameleon' Vine Looks Like Whatever Tree It Climbs". Popular Science. Archived from the original on December 9, 2023. Retrieved December 9, 2023.
  11. ↑ "ScienceShot: 'Chameleon' Vine Discovered in Chile". Science | AAAS. April 24, 2014. Archived from the original on October 31, 2021. Retrieved June 2, 2018.
  12. ↑ Puiu, Tibi (April 25, 2014). "The Chameleon vine: the only plant that morphs host plants near it". ZME Science. Archived from the original on December 9, 2023. Retrieved December 9, 2023.
  13. 1 2 3 4 5 6 7 8 9 10 11 Christenhusz, Maarten J. M. (2012). "738. Boquila Trifoliolata". Curtis's Botanical Magazine. 29 (3): 277–283. doi:10.1111/j.1467-8748.2012.01791.x. ISSN   1355-4905. Archived from the original on December 13, 2023. Retrieved December 13, 2023.
  14. ↑ Valladares, Fernando, Ernesto Gianoli, and Alfredo Saldana. "Climbing plants in a temperate rainforest understorey: searching for high light or coping with deep shade?." Annals of Botany 108.2 (2011): 231-239.
  15. 1 2 3 4 5 Christenhusz, Maarten J. M. (2012). "An Overview of Lardizabalaceae". Curtis's Botanical Magazine. 29 (3): 235–276. doi:10.1111/j.1467-8748.2012.01790.x. ISSN   1355-4905. Archived from the original on November 22, 2022. Retrieved December 14, 2023.
  16. ↑ Vazquez, Miriam Soledad; Rodriguez-Cabal, Mariano A.; Amico, Guillermo C. (March 2022). "The forest gardener: A marsupial with a key seed-dispersing role in the Patagonian temperate forest". Ecological Research. 37 (2): 270–283. Bibcode:2022EcoR...37..270V. doi:10.1111/1440-1703.12289. ISSN   0912-3814. S2CID   245597896. Archived from the original on December 14, 2023. Retrieved December 14, 2023.
  17. ↑ Yirka, Bob; Phys.org. "Researchers discover vine that is able to mimic multiple hosts". phys.org. Retrieved January 10, 2024.
  18. 1 2 3 Wilcox, Christie. "Can Plants See? In the Wake of a Controversial Study, the Answer's Still Unclear". The Scientist Magazine. Archived from the original on December 9, 2023. Retrieved December 9, 2023.
  19. 1 2 3 Gianoli, Ernesto; González-Teuber, Marcia; Vilo, Claudia; Guevara-Araya, María J.; Escobedo, Víctor M. (November 22, 2021). "Endophytic bacterial communities are associated with leaf mimicry in the vine Boquila trifoliolata". Scientific Reports. 11 (1): 22673. Bibcode:2021NatSR..1122673G. doi: 10.1038/s41598-021-02229-8 . ISSN   2045-2322. PMC   8608808 . PMID   34811460.
  20. ↑ Yamashita, Felipe; White, Jacob (2022). "Boquila trifoliolata mimics leaves of an artificial plastic host plant". Plant Signaling & Behavior. 17 (1). Bibcode:2022PlSiB..1777530W. doi:10.1080/15592324.2021.1977530. PMC   8903786 . PMID   34545774.
  21. 1 2 3 Gianoli, Ernesto; González-Teuber, Marcia; Vilo, Claudia; Guevara-Araya, María J.; Escobedo, Víctor M. (November 22, 2021). "Endophytic bacterial communities are associated with leaf mimicry in the vine Boquila trifoliolata". Scientific Reports. 11 (1): 22673. Bibcode:2021NatSR..1122673G. doi: 10.1038/s41598-021-02229-8 . ISSN   2045-2322. PMC   8608808 . PMID   34811460.
  22. ↑ Taylor, Christopher H. (April 1, 2023). "Body size in Batesian mimicry". Evolutionary Ecology. 37 (2): 233–243. Bibcode:2023EvEco..37..233T. doi: 10.1007/s10682-022-10204-6 . ISSN   1573-8477.
  23. ↑ Krulwich, Robert (February 19, 2016). "The Sneaky Life of the World's Most Mysterious Plant". National Geographic. Archived from the original on December 9, 2023. Retrieved December 9, 2023.
  24. 1 2 3 Jones, Benji (November 30, 2022). "The mystery of the mimic plant". Vox. Archived from the original on September 21, 2023. Retrieved December 9, 2023.
  25. 1 2 White, Jacob; Yamashita, Felipe (December 31, 2022). "Boquila trifoliolata mimics leaves of an artificial plastic host plant". Plant Signaling & Behavior. 17 (1). Bibcode:2022PlSiB..1777530W. doi: 10.1080/15592324.2021.1977530 . ISSN   1559-2324. PMC   8903786 . PMID   34545774.
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