treatments-xml/data/03/9C/87/039C87C3FFD6DC3FFF3EE0EFFD60F9C4.xml
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Fossil
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</emphasis>
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<paragraph id="8B8A36D5FFD6DC3FFF3EE0BAFDBAFCE4" blockId="27.[151,1437,506,1652]" pageId="27" pageNumber="726">
In none of the other
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insect orders is the fossil record as poor as in the
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. This is undoubtedly partly due to the special properties of their wings, which not only are particularly fragile because of the paucity (shared with caddisflies) of cross-veins and only moderate sclerotization of the principal longitudinal veins, but, and in particular, because the scale vestiture render them more un-wettable than other insect wings (
<bibRefCitation id="EFA44B24FFD6DC3FFF37E3DAFE21FD04" author="Wagner" box="[158,387,666,692]" pageId="27" pageNumber="740" refString="Wagner, T, Neinhuis, C. &amp; Barthlott, W. (1996) Wettability and contaminability of insect wings as function of their surface sculpures. Acta Zoologica, 77, 213 - 225." type="journal article" year="1996">
Wagner
<emphasis id="B941EAC7FFD6DC3FFEADE3DAFE9BFD03" box="[260,313,666,691]" italics="true" pageId="27" pageNumber="726">et al</emphasis>
. 1996
</bibRefCitation>
)—a factor that evidently impedes their fossilisation in lacustrine (or near-shore marine) sediments. No known fossil
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represent organisation
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that differ markedly from extant ones, and fossils have so far contributed little, if anything, to the understanding of phylogenetic interrelationships within the order. Nevertheless, fossils provide the only direct means for establishing the minimum ages of individual evolutionary lineages.
</paragraph>
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<paragraph id="8B8A36D5FFD6DC3FFF18E222FD60F9C4" blockId="27.[151,1437,506,1652]" pageId="27" pageNumber="726">
The fossil record of
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(as of other insects) was recently reviewed by
<bibRefCitation id="EFA44B24FFD6DC3FFBCEE222FA3AFCCC" author="Grimaldi" box="[1127,1432,866,892]" pageId="27" pageNumber="736" refString="Grimaldi, D. &amp; Engel, M. S. (2005) Evolution of the insects. Cambridge University Press, Cambridge, New York etc., xv + 755 pp." type="book" year="2005">Grimaldi &amp; Engel (2005)</bibRefCitation>
, whose excellent account is recommended strongly. The oldest fossil currently believed to belong to the
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is
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<emphasis id="B941EAC7FFD6DC3FFEBBE2F2FDA3FC7B" box="[274,513,946,971]" italics="true" pageId="27" pageNumber="726">Archaeolepis manae</emphasis>
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from the Lower Jurassic (ca 190 MYO) and according to Grimaldi &amp; Engel a recent re-examination of the specimen has given additional support to its ordinal placement. A small number of moths are known from younger Jurassic strata, but the first lepidopteran fossil that can with any certainty be assigned to a known family lineage (viz.,
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) is from the Lower Cretaceous. The existence of Glossata also in the lower Cretaceous is documented by a larval fossil with a distinct spinneret, while reexamination of an alleged adult glossatan moth from the upper Jurassic failed to confirm the presence of a proboscis. Indeed the presence of glossatans of this age would be unexpected, if angiosperm feeding evolved in the stem lineage of
<taxonomicName id="4C354D56FFD6DC3FFE36E58AFDD3FB54" box="[415,625,1226,1252]" class="Insecta" family="Heterobathmiidae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="726" phylum="Arthropoda" rank="family">Heterobathmiidae</taxonomicName>
+ Glossata. Following
<bibRefCitation id="EFA44B24FFD6DC3FFC2EE58AFB05FB54" box="[903,1191,1226,1252]" pageId="27" pageNumber="738" refString="Labandeira, C. C., Dilcher, D. L., Davis, D. R &amp; Wagner, D. L. (1994) Ninety-seven million years of angiosperm-insect association: Paleobiological insights into the meaning of coevolution. Proceedings of the National Academy of Science, 91, 12278 - 12282." type="journal article">
Labandeira
<emphasis id="B941EAC7FFD6DC3FFBBAE58BFBECFB54" box="[1043,1102,1227,1252]" italics="true" pageId="27" pageNumber="726">et al.</emphasis>
(1994)
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, leaf mines from the Mid Cretaceous Dakota formation attributed to a member of Gracillariidae-Phyllocnistinae have been considered to demarcate the known minimum age of the Ditrysia. Nevertheless, Grimaldi &amp; Engel rightly caution about the uncertainty inherent in identifying leaf miners. In any case there is little doubt that the main radiation of the
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followed the main radiation of angiosperms in the Cretaceous. The point has repeatedly been made that since bat predation is probably the principal selective force behind the evolution of tympanal organs in nocturnal
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, all the tympanate moths lineages (including such species-rich lineages as pyraloids, geometroids and noctuoids) cannot have predated the origin of bats whose currently known minimum age is Early Tertiary. The minimum age for the butterfly families
<taxonomicName id="4C354D56FFD6DC3FFBCCE74AFB67F994" box="[1125,1221,1546,1572]" class="Insecta" family="Pieridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="726" phylum="Arthropoda" rank="family">Pieridae</taxonomicName>
and
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as estimated from molecular evolution is more than 70 MYO, hence the butterflies as a whole must be somewhat older (
<bibRefCitation id="EFA44B24FFD6DC3FFE8BE71AFE51F9C4" author="Braby" box="[290,499,1626,1652]" pageId="27" pageNumber="734" refString="Braby, M. F., Villa, R. &amp; Pierce, N. E. (2006) Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography. Zoological Journal of the Linnean Society, 147, 239 - 275." type="journal article" year="2006">
Braby
<emphasis id="B941EAC7FFD6DC3FFEDAE71BFE0CF9C4" box="[371,430,1627,1652]" italics="true" pageId="27" pageNumber="726">et al.</emphasis>
2006
</bibRefCitation>
,
<bibRefCitation id="EFA44B24FFD6DC3FFE57E71AFD14F9C4" author="Wahlberg" box="[510,694,1626,1652]" pageId="27" pageNumber="740" refString="Wahlberg, N. (2006) That awkward age for butterflies: Insight from the age of the butterfly subfamily Nymphalinae (Lepidoptera: Nymphalidae). Systematic Biology, 55, 703 - 714." type="journal article" year="2006">Wahlberg 2006</bibRefCitation>
).
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