Entomophily

Last updated March 11, 2024

Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, sometimes with conspicuous patterns (honey guides) leading to rewards of pollen and nectar; they may also have an attractive scent which in some cases mimics insect pheromones. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, and in some species also pollen baskets on their hind legs. This required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups. Both the size and the density of a population are known to affect pollination and subsequent reproductive performance.^[1]

Coevolution

History

The insect-pollinated flowers of angiosperms use a combination of cues such as bright colours and streaked patterns to advertise themselves to insects. Flowers-of-Israel-ver006.jpg — The insect-pollinated flowers of angiosperms use a combination of cues such as bright colours and streaked patterns to advertise themselves to insects.

The early spermatophytes (seed plants) were largely dependent on the wind to carry their pollen from one plant to another. Prior to the appearance of flowering plants some gymnosperms, such as Bennettitales, developed flower-like structures that were likely insect pollinated. Insects pollination for gymnosperms likely originated in the Permian period.^[2] Candidates for pollinators include extinct long proboscis insect groups, including Aneuretopsychid, Mesopsychid and Pseudopolycentropodid scorpionflies,^[3] Kalligrammatid ^[4]^[5] and Paradoxosisyrine ^[6] lacewings and Zhangsolvid flies,^[7] as well as some extant families that specialised on gymnosperms before switching to angiosperms, including Nemestrinid, Tabanid and Acrocerid flies.^[8] Living cycads have mutualistic relationships with specific insect species (typically beetles) which pollinate them. Such relationships extend back to at least the late Mesozoic, with both oedemerid beetles (which today are exclusively found on flowering plants)^[2] and boganiid beetles^[9] (which still pollinate cycads today) from the Cretaceous being found with preserved cycad pollen. Angiosperms (flowering plants) first appeared during the Early Cretaceous, and during the angiosperm radiation from 125 to 90 Ma, would displace many of the gymnosperm lineages and cause the extinction of many of their pollinators, while some would transition to angiosperms and some new families would form pollination associations with angiosperms.^[2] Traits such as sapromyophily (emitting the odour of carrion to attract flies) have evolved independently in several unrelated angiosperm families.^[10]

The plant's needs

Wind and water pollination require the production of vast quantities of pollen because of the chancy nature of its deposition. If they are not to be reliant on the wind or water (for aquatic species), plants need pollinators to move their pollen grains from one plant to another. They particularly need pollinators to consistently choose flowers of the same species, so they have evolved different lures to encourage specific pollinators to maintain fidelity to the same species. The attractions offered are mainly nectar, pollen, fragrances and oils. The ideal pollinating insect is hairy (so that pollen adheres to it), and spends time exploring the flower so that it comes into contact with the reproductive structures.^[11]

Mechanisms

Many insects are pollinators, particularly bees, Lepidoptera (butterflies and moths), wasps, flies, ants and beetles.^[11] On the other hand, some plants are generalists, being pollinated by insects in several orders.^[12] Entomophilous plant species have frequently evolved mechanisms to make themselves more appealing to insects, e.g., brightly coloured or scented flowers, nectar, or appealing shapes and patterns. Pollen grains of entomophilous plants are generally larger than the fine pollens of anemophilous (wind-pollinated) plants, which has to be produced in much larger quantities because such a high proportion is wasted. This is energetically costly, but in contrast, entomophilous plants have to bear the energetic costs of producing nectar.^[13]

Hummingbird moth on Clarkia GardenSphinx-20April15.jpg — Hummingbird moth on *Clarkia*

Butterflies and moths have hairy bodies and long proboscides which can probe deep into tubular flowers. Butterflies mostly fly by day and are particularly attracted to pink, mauve and purple flowers. The flowers are often large and scented, and the stamens are so-positioned that pollen is deposited on the insects while they feed on the nectar. Moths are mostly nocturnal and are attracted by night-blooming plants. The flowers of these are often tubular, pale in colour and fragrant only at night. Hawkmoths tend to visit larger flowers and hover as they feed; they transfer pollen by means of the proboscis. Other moths land on the usually smaller flowers, which may be aggregated into flowerheads. Their energetic needs are not so great as those of hawkmoths and they are offered smaller quantities of nectar.^[14]

Inflorescences pollinated by beetles tend to be flat with open corollas or small flowers clustered in a head with multiple, projecting anthers that shed pollen readily.^[11] The flowers are often green or pale-coloured, and heavily scented, often with fruity or spicy aromas, but sometimes with odours of decaying organic matter. Some, like the giant water lily, include traps designed to retain the beetles in contact with the reproductive parts for longer periods.^[15]

Unspecialised flies with short proboscides are found visiting primitive flowers with readily accessible nectar. More specialised flies like syrphids and Tabanids can visit more advanced blooms, but their purpose is to nourish themselves, and any transfer of pollen from one flower to another happens haphazardly. The small size of many flies is often made up for by their abundance, however they are unreliable pollinators as they may bear incompatible pollen, and lack of suitable breeding habitats may limit their activities. Some Pterostylis orchids are pollinated by midges unique to each species. Due to mutual specialisation, pollinators are highly dependent on floral diversity. Therefore, losses in plant diversity, such as those carried on by increasing land use, may be linked to extinctions of pollinators.^[16] A decline, for whatever reason, to one side of this partnership can be catastrophic for the other.^[17]

Flowers pollinated by bees and wasps vary in shape, colour and size. Yellow or blue plants are often visited, and flowers may have ultra-violet nectar guides, that help the insect to find the nectary. Some flowers, like sage or pea, have lower lips that will only open when sufficiently heavy insects, such as bees, land on them. With the lip depressed, the anthers may bow down to deposit pollen on the insect's back. Other flowers, like tomato, may only liberate their pollen by buzz pollination, a technique in which a bumblebee will cling on to a flower while vibrating its flight muscles, and this dislodges the pollen. Because bees care for their brood, they need to collect more food than just to maintain themselves, and therefore are important pollinators.^[17] Other bees are nectar thieves and bite their way through the corolla in order to raid the nectary, in the process bypassing the reproductive structures.^[11]

Ants are not well adapted to pollination but they have been shown to perform this function in Polygonum cascadense and in certain desert plants with small blossoms near the ground with little fragrance or visual attraction, small quantities of nectar and limited quantities of sticky pollen.^[17]

Plant-insect pairings

Some plant species co-evolved with a particular pollinator species, such as the bee orchid. The species is almost exclusively self-pollinating in its northern ranges, but is pollinated by the solitary bee Eucera in the Mediterranean area. The plant attracts these insects by producing a scent that mimics the scent of the female bee. In addition, the lip acts as a decoy, as the male bee confuses it with a female that is visiting a pink flower. Pollen transfer occurs during the ensuing pseudocopulation.^[18]

Figs in the genus Ficus have a mutualistic arrangement with certain tiny agaonid wasps. In the common fig, the inflorescence is a syconium, formed by an enlarged, fleshy, hollow receptacle with multiple ovaries on the inner surface. A female wasp enters through a narrow aperture, fertilizes these pistillate flowers, and lays its eggs in some ovaries, with galls being formed by the developing larvae. In due course, staminate flowers develop inside the syconium. Wingless male wasps hatch and mate with females in the galls before tunnelling their way out of the developing fruit. The winged females, now laden with pollen, follow, flying off to find other receptive syconia at the right stage of development. Most species of fig have their own unique commensal species of wasp.^[19]

Etymology

The word is artificially derived from the Greek: εντομο-, entomo-^[20] "cut in pieces, segmented", hence "insect"; and φίλη, phile, "loved".

Taxonomic range

Wind pollination is the reproductive strategy adopted by the grasses, sedges, rushes and catkin-bearing plants. Other flowering plants are mostly pollinated by insects (or birds or bats), which seems to be the primitive state, and some plants have secondarily developed wind pollination. Some plants that are wind pollinated have vestigial nectaries, and other plants like common heather that are regularly pollinated by insects, produce clouds of pollen and some wind pollination is inevitable. The hoary plantain is primarily wind pollinated, but is also visited by insects which pollinate it.^[13] In general, showy, colourful, fragrant flowers like sunflowers, orchids and Buddleja are insect pollinated. The only entomophilous plants that are not seed plants are the dung-mosses of the family Splachnaceae.^[21]

Related Research Articles

In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. It often contains the nectaries of the plant. It is present in many plant families, although varies in structural dimensions and appearance. This differentiation between the hypanthium in particular species is useful for identification. Some geometric forms are obconic shapes as in toyon, whereas some are saucer-shaped as in Mitella caulescens.

A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains.

Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the corolla. Petals are usually accompanied by another set of modified leaves called sepals, that collectively form the calyx and lie just beneath the corolla. The calyx and the corolla together make up the perianth, the non-reproductive portion of a flower. When the petals and sepals of a flower are difficult to distinguish, they are collectively called tepals. Examples of plants in which the term tepal is appropriate include genera such as Aloe and Tulipa. Conversely, genera such as Rosa and Phaseolus have well-distinguished sepals and petals. When the undifferentiated tepals resemble petals, they are referred to as "petaloid", as in petaloid monocots, orders of monocots with brightly colored tepals. Since they include Liliales, an alternative name is lilioid monocots.

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 as gene-culture coevolution.

Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, for example beetles or butterflies; birds, and bats; water; wind; and even plants themselves. Pollinating animals travel from plant to plant carrying pollen on their bodies in a vital interaction that allows the transfer of genetic material critical to the reproductive system of most flowering plants. When self-pollination occurs within a closed flower. Pollination often occurs within a species. When pollination occurs between species, it can produce hybrid offspring in nature and in plant breeding work.

<span class="mw-page-title-main">Gymnosperm</span> Clade of non-flowering, naked-seeded vascular plants

The gymnosperms are a group of seed-producing plants that includes conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, Ginkgo. Gymnosperm lifecycles involve alternation of generations. They have a dominant diploid sporophyte phase and a reduced haploid gametophyte phase which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.

Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales were amongst the most common seed plants of the Mesozoic, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.

Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous plants are oaks, pecans, pistachios, sweet chestnuts, alders and members of the family Juglandaceae. Approximately 12% of plants across the globe are pollinated by anemophily, including cereal crops like rice and corn and other prominent crop plants like wheat, rye, barley, and oats. In addition, many pines, spruces, and firs are wind-pollinated.

Zoophily, or zoogamy, is a form of pollination whereby pollen is transferred by animals, usually by invertebrates but in some cases vertebrates, particularly birds and bats, but also by other animals. Zoophilous species frequently have evolved mechanisms to make themselves more appealing to the particular type of pollinator, e.g. brightly colored or scented flowers, nectar, and appealing shapes and patterns. These plant-animal relationships are often mutually beneficial because of the food source provided in exchange for pollination.

<span class="mw-page-title-main">Nectar</span> Sugar-rich liquid produced by many flowering plants, that attracts pollinators and insects

Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, which in turn provide herbivore protection. Common nectar-consuming pollinators include mosquitoes, hoverflies, wasps, bees, butterflies and moths, hummingbirds, honeyeaters and bats. Nectar plays a crucial role in the foraging economics and evolution of nectar-eating species; for example, nectar foraging behavior is largely responsible for the divergent evolution of the African honey bee, A. m. scutellata and the western honey bee.

A flower, also known as a bloom or blossom, is the reproductive structure found in flowering plants. Flowers consist of a combination of vegetative organs – sepals that enclose and protect the developing flower, petals that attract pollinators, and reproductive organs that produce gametophytes, which in flowering plants produce gametes. The male gametophytes, which produce sperm, are enclosed within pollen grains produced in the anthers. The female gametophytes are contained within the ovules produced in the carpels.

The family Oedemeridae is a cosmopolitan group of beetles commonly known as false blister beetles, though some recent authors have coined the name pollen-feeding beetles. There are some 100 genera and 1,500 species in the family, mostly associated with rotting wood as larvae, though adults are quite common on flowers. The family was erected by Pierre André Latreille in 1810.

In zoology, a palynivore /pəˈlɪnəvɔːɹ/, meaning "pollen eater" is an herbivorous animal which selectively eats the nutrient-rich pollen produced by angiosperms and gymnosperms. Most true palynivores are insects or mites. The category in its strictest application includes most bees, and a few kinds of wasps, as pollen is often the only solid food consumed by all life stages in these insects. However, the category can be extended to include more diverse species. For example, palynivorous mites and thrips typically feed on the liquid content of the pollen grains without actually consuming the exine, or the solid portion of the grain. Additionally, the list is expanded greatly if one takes into consideration species where either the larval or adult stage feeds on pollen, but not both. There are other wasps which are in this category, as well as many beetles, flies, butterflies, and moths. One such example of a bee species that only consumes pollen in its larval stage is the Apis mellifera carnica. There is a vast array of insects that will feed opportunistically on pollen, as will various birds, orb-weaving spiders and other nectarivores.

<span class="mw-page-title-main">Pollination syndrome</span> Flower traits that attract pollinators

Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different pollen vectors, which can be abiotic or biotic, such as birds, bees, flies, and so forth through a process called pollinator-mediated selection. These traits include flower shape, size, colour, odour, reward type and amount, nectar composition, timing of flowering, etc. For example, tubular red flowers with copious nectar often attract birds; foul smelling flowers attract carrion flies or beetles, etc.

Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.

Anthecology, or pollination biology, is the study of pollination as well as the relationships between flowers and their pollinators. Floral biology is a bigger field that includes these studies. Most flowering plants, or angiosperms, are pollinated by animals, and especially by insects. The major flower-frequenting insect taxa include beetles, flies, wasps, bees, ants, thrips, butterflies, and moths. Insects carry out pollination when visiting flowers to obtain nectar or pollen, to prey on other species, or when pseudo-copulating with insect-mimicking flowers such as orchids. Pollination-related interactions between plants and insects are considered mutualistic, and the relationships between plants and their pollinators have likely led to increased diversity of both angiosperms and the animals that pollinate them.

Pollination traps or trap-flowers are plant flower structures that aid the trapping of insects, mainly flies, so as to enhance their effectiveness in pollination. The structures of pollination traps can include deep tubular corollas with downward pointing hairs, slippery surfaces, adhesive liquid, attractants, flower closing and other mechanisms.

The monocots are one of the two major groups of flowering plants, the other being the dicots. In order to reproduce they utilize various strategies such as employing forms of asexual reproduction, restricting which individuals they are sexually compatible with, or influencing how they are pollinated. Nearly all reproductive strategies that evolved in the dicots have independently evolved in monocots as well. Despite these similarities and their close relatedness, monocots and dicots have distinct traits in their reproductive biologies.

The pollination of orchids is a complex chapter in the biology of this family of plants that are distinguished by the complexity of their flowers and by intricate ecological interactions with their pollinator agents. It has captured the attention of numerous scientists over time, including Charles Darwin, father of the theory of evolution by natural selection. Darwin published in 1862 the first observations of the fundamental role of insects in orchid pollination, in his book The Fertilization of Orchids. Darwin stated that the varied stratagems orchids use to attract their pollinators transcend the imagination of any human being.

<span class="mw-page-title-main">UV coloration in flowers</span> Natural phenomenon

UV coloration is a natural phenomenon that leads to unique interactions between organisms that have evolved the ability to perceive these wavelengths of light. It serves as one method to attract pollinators to the flower along with scent, shape, and nectar quality. Flowers are known for their range of visible colors that humans can see with their eyes and observe an array of different shades and patterns. The naked eye cannot see the ultraviolet coloration many flowers employ to bring attention to themselves. By either reflecting or absorbing UV light waves, flowers are able to communicate with pollinators. This allows plants that may require an animal pollinator to stand out from other flowers or distinguish where their flowers are in a muddied background of other plant parts. For the plant, it is important to share and receive pollen so they can reproduce, maintain their ecological role, and guide the evolutionary history of the population.

References

↑ Mustajärvi, Kaisa; Siikamäki, Pirkko; Rytkönen, Saara; Lammi, Antti (2001). "Consequences of plant population size and density for plant-pollinator interactions and plant performance: Plant-pollinator interactions". Journal of Ecology. 89 (1): 80–87. doi: 10.1046/j.1365-2745.2001.00521.x .
1 2 3 Peris, David; Pérez-de la Fuente, Ricardo; Peñalver, Enrique; Delclòs, Xavier; Barrón, Eduardo; Labandeira, Conrad C. (March 2017). "False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance". Current Biology. 27 (6): 897–904. doi: 10.1016/j.cub.2017.02.009 . hdl: 2445/163381 . ISSN 0960-9822. PMID 28262492. S2CID 3967504.
↑ Zhao, Xiangdong; Wang, Bo; Bashkuev, Alexey S.; Aria, Cédric; Zhang, Qingqing; Zhang, Haichun; et al. (March 2020). "Mouthpart homologies and life habits of Mesozoic long-proboscid scorpionflies". Science Advances. 6 (10): eaay1259. Bibcode:2020SciA....6.1259Z. doi:10.1126/sciadv.aay1259. PMC 7056314 . PMID 32181343.
↑ Labandeira, Conrad C.; Yang, Qiang; Santiago-Blay, Jorge A.; Hotton, Carol L.; Monteiro, Antónia; Wang, Yong-Jie; Goreva, Yulia; Shih, ChungKun; Siljeström, Sandra; Rose, Tim R.; Dilcher, David L. (2016-02-10). "The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies". Proceedings of the Royal Society B: Biological Sciences. 283 (1824): 20152893. doi:10.1098/rspb.2015.2893. ISSN 0962-8452. PMC 4760178 . PMID 26842570.
↑ Liu, Qing; Lu, Xiumei; Zhang, Qingqing; Chen, Jun; Zheng, Xiaoting; Zhang, Weiwei; Liu, Xingyue; Wang, Bo (2018-09-17). "High niche diversity in Mesozoic pollinating lacewings". Nature Communications. 9 (1): 3793. Bibcode:2018NatCo...9.3793L. doi: 10.1038/s41467-018-06120-5 . ISSN 2041-1723. PMC 6141599 . PMID 30224679.
↑ Khramov, Alexander V.; Yan, Evgeny; Kopylov, Dmitry S. (December 2019). "Nature's failed experiment: Long-proboscid Neuroptera (Sisyridae: Paradoxosisyrinae) from Upper Cretaceous amber of northern Myanmar". Cretaceous Research. 104: 104180. doi:10.1016/j.cretres.2019.07.010. S2CID 199111088.
↑ Peñalver, Enrique; Arillo, Antonio; Pérez-de la Fuente, Ricardo; Riccio, Mark L.; Delclòs, Xavier; Barrón, Eduardo; Grimaldi, David A. (July 2015). "Long-Proboscid Flies as Pollinators of Cretaceous Gymnosperms". Current Biology. 25 (14): 1917–1923. doi: 10.1016/j.cub.2015.05.062 . PMID 26166781. S2CID 13022302.
↑ Khramov, Alexander V.; Lukashevich, Elena D. (2019). "A Jurassic dipteran pollinator with an extremely long proboscis". Gondwana Research. 71: 210–215. Bibcode:2019GondR..71..210K. doi:10.1016/j.gr.2019.02.004. S2CID 134847380.
↑ Cai, Chenyang; Escalona, Hermes E.; Li, Liqin; Yin, Ziwei; Huang, Diying; Engel, Michael S. (September 2018). "Beetle Pollination of Cycads in the Mesozoic". Current Biology. 28 (17): 2806–2812.e1. doi: 10.1016/j.cub.2018.06.036 . PMID 30122529. S2CID 52038878.
↑ McGhee, George R. (2011). Convergent Evolution: Limited Forms Most Beautiful. MIT Press. pp. 118–120. ISBN 978-0-262-01642-1.
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↑ Köhler, Andreas; Sühs, Rafael Barbizan; Somavilla, Alexandre (2010-11-11). "Entomofauna associated to the floration of Schinus terebinthifolius Raddi (Anacardiaceae) in the Rio Grande do Sul State, Brazil = Entomofauna associada à floração de Schinus terebinthifolius Raddi (Anacardiaceae) no Estado do Rio Grande do Sul, Brasil | Somavilla |". Bioscience Journal. 26 (6). Retrieved 18 April 2014.
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↑ Oliveira, P.E.; Gibbs, P.E.; Barbosa, A.A. (2004). "Moth pollination of woody species in the Cerrados of Central Brazil: a case of so much owed to so few?". Plant Systematics and Evolution. 245 (1–2): 41–54. doi:10.1007/s00606-003-0120-0. S2CID 21936259.
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↑ Fründ, Jochen; Linsenmair, Karl Eduard; Blüthgen, Nico (2010-09-14). "Pollinator diversity and specialization in relation to flower diversity". Oikos. 119 (10): 1581–1590. doi:10.1111/j.1600-0706.2010.18450.x. ISSN 0030-1299.
1 2 3 Faegri, K.; Van Der Pijl, L. (2013). Principles of Pollination Ecology. Elsevier. pp. 102–110. ISBN 978-1-4832-9303-5.
↑ Fenster, Charles B.; Marten-Rodriguez, Silvana (2007). "Reproductive Assurance And The Evolution Of Pollination Specialization" (PDF). International Journal of Plant Sciences. 168 (2): 215–228. doi:10.1086/509647. S2CID 1890083.
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↑ Goffinet, Bernard; Shaw, A. Jonathan; Cox, Cymon J. (2004). "Phylogenetic inferences in the dung-moss family Splachnaceae from analyses of cpDNA sequence data and implications for the evolution of entomophily". American Journal of Botany . 91 (5): 748–759. doi:10.3732/ajb.91.5.748. PMID 21653429.

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[1] Mustajärvi, Kaisa; Siikamäki, Pirkko; Rytkönen, Saara; Lammi, Antti (2001). "Consequences of plant population size and density for plant-pollinator interactions and plant performance: Plant-pollinator interactions". Journal of Ecology. 89 (1): 80–87. doi: 10.1046/j.1365-2745.2001.00521.x .

[:0-2] 1 2 3 Peris, David; Pérez-de la Fuente, Ricardo; Peñalver, Enrique; Delclòs, Xavier; Barrón, Eduardo; Labandeira, Conrad C. (March 2017). "False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance". Current Biology. 27 (6): 897–904. doi: 10.1016/j.cub.2017.02.009 . hdl: 2445/163381 . ISSN 0960-9822. PMID 28262492. S2CID 3967504.

[3] Zhao, Xiangdong; Wang, Bo; Bashkuev, Alexey S.; Aria, Cédric; Zhang, Qingqing; Zhang, Haichun; et al. (March 2020). "Mouthpart homologies and life habits of Mesozoic long-proboscid scorpionflies". Science Advances. 6 (10): eaay1259. Bibcode:2020SciA....6.1259Z. doi:10.1126/sciadv.aay1259. PMC 7056314 . PMID 32181343.

[4] Labandeira, Conrad C.; Yang, Qiang; Santiago-Blay, Jorge A.; Hotton, Carol L.; Monteiro, Antónia; Wang, Yong-Jie; Goreva, Yulia; Shih, ChungKun; Siljeström, Sandra; Rose, Tim R.; Dilcher, David L. (2016-02-10). "The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies". Proceedings of the Royal Society B: Biological Sciences. 283 (1824): 20152893. doi:10.1098/rspb.2015.2893. ISSN 0962-8452. PMC 4760178 . PMID 26842570.

[5] Liu, Qing; Lu, Xiumei; Zhang, Qingqing; Chen, Jun; Zheng, Xiaoting; Zhang, Weiwei; Liu, Xingyue; Wang, Bo (2018-09-17). "High niche diversity in Mesozoic pollinating lacewings". Nature Communications. 9 (1): 3793. Bibcode:2018NatCo...9.3793L. doi: 10.1038/s41467-018-06120-5 . ISSN 2041-1723. PMC 6141599 . PMID 30224679.

[6] Khramov, Alexander V.; Yan, Evgeny; Kopylov, Dmitry S. (December 2019). "Nature's failed experiment: Long-proboscid Neuroptera (Sisyridae: Paradoxosisyrinae) from Upper Cretaceous amber of northern Myanmar". Cretaceous Research. 104: 104180. doi:10.1016/j.cretres.2019.07.010. S2CID 199111088.

[7] Peñalver, Enrique; Arillo, Antonio; Pérez-de la Fuente, Ricardo; Riccio, Mark L.; Delclòs, Xavier; Barrón, Eduardo; Grimaldi, David A. (July 2015). "Long-Proboscid Flies as Pollinators of Cretaceous Gymnosperms". Current Biology. 25 (14): 1917–1923. doi: 10.1016/j.cub.2015.05.062 . PMID 26166781. S2CID 13022302.

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