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Aglossata Heterobathmiina Glossata 29 Zeugloptera

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Aglossata Heterobathmiina Glossata 29 Zeugloptera 1 Noctuoidea (owlet moths) 1 Geometroidea Drepanoidea (hook tip moths) Papilionoidea (butteries) 2 Calliduloidea 2 Cimelioidea (gold moths) Bombycoidea (silk moths, sphinx moths) 3 Lasiocampoidea (lappet moths) Mimallonoidea (sack bearer moths) 3 Thyridoidea (picture-winged leaf moths) Pyraloidea 4 Hyblaeoidea (teak moths) Copromorphoidea (fruitworm moths) Immoidea 4 Whalleyanoidea Pterophoroidea (plume moths) Alucitoidea (many-plumed moths) Ditrysia Epermenioidea 5 Schreckensteinioidea Urodoidea (false burnet moths) Tortricoidea (leafroller moths) 5 Choreutoidea Sesioidea (clearwing moths) GLOSSATA 6 Cossoidea (carpenter moths) Zygaenoidea (burnet moths, forester moths) 6 Galacticoidea Symaethistoidea Gelechioidea (case-bearers, twirler moths) 7 Yponomeutoidea (ermine moths) 7 Gracillarioidea (leaf miners) Tineoidea (cloth moths) 8 Tischerioidea 8 Palaephatoidea Adeloidea 9 Andesianoidea Nepticuloidea Hepialoidea (ghost moths, swift moths) 10 9 Mnesarchaeoidea Neopseustoidea Lophocoronoidea Acanthopteroctetoidea 10 MYA Eriocranioidea Heterobathmioidea HETEROBATHMIINA c. 160 Agathiphagoidea AGLOSSATA 29 Micropterigoidea ZEUGLOPTERA Tree of Lepidopterans showing the phylogenetic relationships among the most significant groups (45 taxonomic superfamilies). Colored boxes indicate high taxonomic ranks. Branches with thick lines indicate robust clades, and branches with thin lines indicate less-supported clades. The number in the green circle indicates the chapter in which lepidopterans are also included. The orange circle marks the most internal node and its age. Photographs illustrate principal clades; boxed numbers associate photographs with clades. © 2014 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 36_TOL1e_36.indd 428 5/8/14 11:13 AM Lepidopterans Butterflies and Moths Roger Vila 36 SUMMARY Both professional researchers and aficionados, motivated by the appeal and popularity of Lepidoptera (butterflies and moths), have compiled a huge amount of knowl- edge on their biology, taxonomy, and distribution over the centuries, comparable only to the amount of information we have on vertebrates. However, it is only recently that a clear picture of the branch of the tree of life belonging to these insects has begun to emerge. The huge diversity of lepidopterans poses considerable difficulties for studying their evolu- tion: with 170,000 species described, Lepidoptera represents the largest lineage of herbiv- orous organisms and one of the four most diverse orders on the planet. The limited fossil record has been another obstacle, both for understanding the evolution of morphological characters and for calibrating molecular clocks. The results of phylogenetic studies based on nucleotide sequences generally support hypotheses based on morphological charac- ters. Indeed, both methods are advancing in parallel, and as more characters are added, the results are generally congruent. The taxonomy is gradually being adapted to phyloge- netic results as they are consolidated and confirmed in numerous independent studies. To- day, the most basal relationships are quite well resolved, as is the general structure of the tree. Along with fossil information, it tells the story of lepidopterans: their emergence in the Triassic or Early Jurassic from a common ancestor resembling the caddisfly (Trichoptera), the evolution from mandibulates to microlepidoptera with a proboscis, and the progressive acquisition of other specialized characters throughout one of the lineages, leading to the huge radiations of Ditrysia, which include the large moths and butterflies. Most of the rela- tionships within the Ditrysia have yet to be satisfactorily resolved, due largely to this group’s extremely rapid diversification. Projects under way include research on the phylogeny of the order based on morphological data and a large number of nuclear genes, the use of phylogenomics to resolve the most difficult cases, and investigation of multiple phylogenies of superfamilies and lower taxa. These promise exciting results that will clear up many ob- scure points in lepidopteran phylogeny and evolution. With some 170,000 species described, lepidopterans, the most extensive for any group of such great biodi- along with coleopterans, hymenopterans, and dipter- versity. Their evolutionary success is based at least in ans, constitute one of the most diverse orders on the part on their holometabolism, which allows for a high planet. Their beauty has captured the interest of scien- degree of specialization in their different life stages tists and aficionados alike, and the information com- (see Figure 36.1). Another key factor is their special- piled on their biology, taxonomy, and distribution is ized mouthpart (proboscis or haustellum), which al- What is a lepidopteran? A lepidopteran (from the Greek lepis [scale] and contain pigments and microstructures with optical pteron [wing]) is a holometabolous (undergoing properties, giving their color patterns enormous complete metamorphosis) insect whose larvae, plasticity. The order Lepidoptera, one of the most called caterpillars, are herbivorous, with rare diverse orders on the planet, is the sister clade exceptions. It has six true legs and multiple prolegs of the trichopterans or caddisflies (Trichoptera), with diminutive hooks on them. The adults develop and together these clades form the superorder two pairs of wings densely covered in scales that Amphiesmenoptera. © 2014 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 36_TOL1e_36.indd 429 5/8/14 11:13 AM 430 CHAPTER 36 • LEPIDOPTERANS (A) (B) insects. Although most species have this chromosome number, there are notable exceptions. For example, the butterfly family Lycaenidae has a modal number of 23–24. In some clades, the variability in chromosome (C) number is huge. For example, the subgenus Agrodiae- tus (Papilionoidea, Lycaenidae) contains species with chromosome numbers ranging from 10 to 134. Poly- ommatus atlanticus (Figure 36.2), also a lycaenid, is the metazoan with the highest chromosome number: its haploid number is around 223. Leptidea sinapis (Papil- ionoidea, Pieridae) boasts another record: it exhibits a cline (continuous gradient) in number of chromosomes from n = 53 on the Iberian Peninsula to n = 28 in Central Figure 36.1 Postembryonic stages of a holometabolous insect, the lepidopteran Melitaea didyma (Papilionoidea: Asia. Surprisingly, the extremes have been observed to Nymphalidae). (A) Larva, (B) pupa, and (C) adult. maintain a notable level of fertility when crossed. An- other curiosity of lepidopteran cytogenetics is that they Figure 36.1 have achiasmatic oogenesis; in other words, there is lows them to sip nectar from flowering plants (angio- no meiotic recombination of maternal genetic mate- sperms). Lepidopterans are the largest lineage of phy- rial. Knowledge of this group’s chromosomal evolu- tophagous organisms in existence, as their larvae feed tion, the type of rearrangements, and the reasons for mostly on plants, especially angiosperms. These two their occurrence is practically nonexistent. It seems that groups have therefore evolved in tandem, and it is not karyotype changes occur as a result of fusion and frag- surprising that the explosive radiation of angiosperms mentation, given that the total amount of genetic mate- (see Chapter 12, Angiosperms) coincides with that of rial apparently remains constant (see also Chapter 12, butterflies and moths, as well as that of other pollinat- Angiosperms). The DNA content (C-value) has been ing insects like bees (see Chapter 33, Hymenopterans). determined for 60 species of lepidopterans, with an av- Another very important characteristic in the basic anat- erage value of 0.66 ± 0.04 picogram (pg), ranging from omy of lepidopterans is the dense covering of scale on a minimum of 0.29 to a maximum of 1.9 pg. their wings. This serves a function in both flight and The most commonly used markers in phylogenet- thermoregulation, and also gives their color patterns ics include mitochondrial genes (cox1, cox2, nad1 and enormous plasticity, a trait with profound implications nad5) and single-copy nuclear genes (CAD, EF-1a, for intra- and inter-specific relationships. H3, wg, etc.), as well as ribosomal regions (mainly the Knowledge of the lepidopteran evolutionary tree subunit 28S), and noncoding regions such as spacer is still only partial (see Tree of Lepidopterans), and un- regions (ITS-2) of the nuclear genome. Mitochondrial til recently was based only on morphological charac- markers have a greater substitution rate than nuclear ters. Studies using modern techniques of phylogenetic markers, and are therefore very helpful for studying reconstruction have appeared more recently, as have the relationship between only slightly divergent taxa, studies based on DNA sequences, or on a combination of morphological and molecular data. This chapter (A) (B) describes the phylogeny and evolution of lepidopter- ans, with special reference to results of the most recent studies. Characteristics of Lepidopteran Genomes Our knowledge of lepidopteran cytogenetics, as well as (C) that of trichopterans with whom they share the char- acteristics described below, is limited. Their sex-de- termination system is ZW (heterogametic females). Their chromosomes are holocentric, appearing almost
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