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Prodoxidae
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==Yucca moths and coevolution== "Yucca moths" have a remarkable biology. They are famous for an old and intimate relationship with ''[[Yucca]]'' plants and are their [[obligate]] [[pollinators]] as well as [[herbivores]].<ref name=Pellmyr-eal-1996>{{cite journal |last1=Pellmyr |first1=Olle |last2=Thompson |first2=John N. |last3=Brown |first3=Johnathan M. |last4=Harrison |first4=Richard G. |date=1996 |title=Evolution of pollination and mutualism in the yucca moth lineage |journal=American Naturalist |volume=148 |issue=5 |pages=827β847 |doi=10.1086/285958 |jstor=2463408 |bibcode=1996ANat..148..827P |s2cid=84816447 }}</ref> Interactions of these organisms range from obligate [[Mutualism (biology)|mutualism]] to [[commensalism]] to outright [[antagonism (phytopathology)|antagonism]]. Their bore holes are a common sight on trunks of such plants as the [[soaptree yucca]]. Two of the three yucca moth genera in particular, ''[[Tegeticula]]'' and ''[[Parategeticula]]'', have an obligate pollination mutualism with yuccas. Yuccas are only pollinated by these moths, and the [[pollinator]] [[larva]]e feed exclusively on yucca [[seed]]s; the female moths use their modified mouthparts to insert the pollen into the stigma of the flowers, after having oviposited in the ovary, where the larvae feed on some (but not all) of the developing ovules. This obligate pollination mutualism is similar to the mutualistic relationship between the [[Pachycereus schottii|senita cactus]] and the [[Upiga|senita moth]].<ref>{{cite journal |last1=Holland |first1=J. Nathaniel |last2=Fleming |first2=Theodore H. |date=1999-09-01 |title=Mutualistic interactions between ''Upiga virescens'' (Pyralidae), a pollinating seed-consumer, and ''Lophocereus schottii'' (Cactaceae) |journal=Ecology |language=en |volume=80 |issue=6 |pages=2074β2084 |doi=10.1890/0012-9658(1999)080[2074:mibuvp]2.0.co;2 |issn=1939-9170 |hdl=1911/21700 |url=https://scholarship.rice.edu/bitstream/1911/21700/1/1999aHolland_Fleming_Ecology.pdf|hdl-access=free }}</ref> Species of the third genus of yucca moths, ''[[Prodoxus]]'', are not engaged in the pollination mutualism, nor do the larvae feed on developing seeds. Their [[Egg (biology)|egg]]s are deposited in [[fruit]]s and leaves, where they eat and grow, not emerging until fully mature. One species of yucca moth, ''[[Tegeticula intermedia]],'' betrays this obligate mutualism by not pollinating the yucca while still laying its eggs on the host plant, cheating the yucca out of any benefits from this relationship.<ref>{{Cite journal|last1=Marr|first1=Deborah L.|last2=Brock|first2=Marcus T.|last3=Pellmyr|first3=Olle|date=2001-08-01|title=Coexistence of mutualists and antagonists: exploring the impact of cheaters on the yucca β yucca moth mutualism|journal=Oecologia|language=en|volume=128|issue=3|pages=454β463|doi=10.1007/s004420100669|pmid=24549915|bibcode=2001Oecol.128..454M|issn=0029-8549|hdl=2022/24397|s2cid=6432027|hdl-access=free}}</ref> [[Coevolution]] is particularly important in evolutionary biology as it demonstrates increased genetic variance between two organisms that have strong interactions, resulting in increased fitness generally for both species. In an effort to further investigate the traits that have evolved as a result of coevolution, Pellmyr and his team utilized a phylogenetic framework to observe the evolution of active pollination and specializing effects of the yucca moths which eventually lead to the loss of nectar in the genus of yucca plants, requiring them to have Prodoxidae moths around to reproduce. The moths in this case, specifically ''Tegeticula'' and ''Parategeticula'', pollinate yucca flower purposefully, and lay their eggs in the flowers. The larvae of the moths rely on yucca seeds as nourishment and this is also cost inflicted on the plants to maintain the mutualism. After setting up a test experiment which involved pairing species of Prodoxidae with different host plants, the results have shown that moths that were able to develop a pollination-type relationship with the new plant species were more successful and would better be able to reproduce than moths that were unable to do so.<ref name=Pellmyr-eal-1996/><ref name=Groman-Pellmyr-2000>{{cite journal |last1=Groman |first1=Joshua D. |last2=Pellmyr |first2=Olle |date=2000 |title=Rapid evolution and specialization following host colonization in a yucca moth |journal=Journal of Evolutionary Biology |volume=13 |issue=2 |pages=223β236 |doi=10.1046/j.1420-9101.2000.00159.x |s2cid=84556390 |doi-access=free }}</ref> Another study takes a look at coevolution as a primary driver of change and diversification in the yucca moth and the [[Joshua tree]], more commonly known as the yucca palm. The researchers tested this hypothesis by setting up a differential selection of two species of yucca moths and two corresponding species of yucca palms which they pollinate. The study showed floral traits involving pollination evolved substantially more rapidly than other flower features. The study then looks at phylogeny and determines that coevolution is the major evolutionary force behind diversification in the yucca palms when pollinated moths were present. The researchers of the Joshua tree show that setting up phylogenetic patterns using maximum likelihood techniques, can be a powerful tool to analyze the divergence in species.<ref>{{cite journal |last1=Godsoe |first1=William |last2=Yoder |first2=Jeremy B. |author2-link=Jeremy Yoder |last3=Smith |first3=Christopher Irwin |last4=Pellmyr |first4=Olle |date=2008-04-04 |title=Coevolution and divergence in the Joshua tree/yucca moth mutualism |journal=The American Naturalist |volume=171 |issue=6 |pages=816β823 |doi=10.1086/587757 |jstor=10.1086/587757 |pmid=18462130 |bibcode=2008ANat..171..816G |s2cid=12410715 }}</ref> Researchers have again tried to demonstrate the absolute minimal level of evolution needed to secure a yucca plant and moth mutualism. The researchers attempt to find an answer as to how integral coevolution was as the driving force behind various adaptions between the yucca moth and plant species. Phylogenetic examination was also used here to reconstruct the trait evolution of the pollinating yucca moths and their non-mutualistic variants. Certain mutualistic traits have predated the yucca moth-plant mutualism, such as larval feeding in the floral ovary; however, it suggests that other key traits linked to pollination were indeed a result of coevolution between the two species. It is integral to reiterate here that key traits such as tentacular appendages which help in pollen collection and pollinating behaviors evolved as a result of coevolution during a mutualism between moths and host plants. After collecting genetic information from dozens of differing Prodoxidae moths, including ones involved in ideal mutualisms such as ''Tegeticula'', and mapping these extracted sequences using the BayesTraits clade forming algorithm, conclusions could be drawn about trait formation that differentiated the monophylum or clade of strict obligate pollinators in the family Prodoxidae from other moths that did not undergo mutualism.<ref name=Yoder-eal-2010>{{cite journal |last1=Yoder |first1=Jeremy B. |author1-link=Jeremy Yoder |last2=Smith |first2=Christopher Irwin |last3=Pellmyr |first3=Olle |date=2010-08-01 |title=How to become a yucca moth: minimal trait evolution needed to establish the obligate pollination mutualism |journal=Biological Journal of the Linnean Society |volume=100 |issue=4 |pages=847β855 |doi=10.1111/j.1095-8312.2010.01478.x |pmid=20730026 |pmc=2922768 }}</ref>
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