Difference between revisions of "AY Honors/Insect/Lepidoptera"

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{{Taxobox
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{{Species id
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| name = [[Butterfly|Butterflies]], [[moth]]s and [[Skipper (butterfly)|skippers]]
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| common_name = Butterflies, Moths and Skippers
 
| image = Parthenos_sylvia_philippensis.jpg
 
| image = Parthenos_sylvia_philippensis.jpg
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| image_width = 250px
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| latin_name = Lepidoptera
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| image_caption = The Clipper ''Parthenos sylvia''
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| order = '''Lepidoptera'''
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| regnum = [[Animal]]ia
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| description =
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| phylum = [[Arthropod]]a
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Lepidopterans  undergo complete metamorphosis, going through a four-stage life cycle:
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| classis = [[Insect]]a
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# egg
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| subclassis = [[Pterygota]]
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# larva/caterpillar  
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| infraclassis = [[Neoptera]]
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# pupa/chrysalis
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| superordo = [[Endopterygota]]
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# imago/adult.
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| ordo = '''Lepidoptera'''
 
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| ordo_authority = [[Carolus Linnaeus|Linnaeus]], [[Systema Naturae|1758]]
 
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| subdivision_ranks = Subdivisions
 
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| subdivision =
 
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See [[Taxonomy of Lepidoptera]] and [[Lepidopteran diversity]].
 
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}}
 
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The order '''Lepidoptera''' is the one of the most speciose [[order (biology)|order]]s in the [[class (biology)|class]] [[Insect]]a and includes the [[butterfly|butterflies]], [[moth]]s and [[skipper (butterfly)|skipper]]s.  Members of the order are referred to as '''lepidopterans'''. A person who collects or studies this order is referred to as a [[lepidopterist]].  This order has more than 180,000 [[species]]<ref>
 
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{{cite web| url = http://www.ucl.ac.uk/taxome/ | title = The Lepidoptera Taxome Project Draft Proposals and Information
 
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| accessdate = 2007-03-05
 
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| publisher = Centre for Ecology and Evolution, University College London
 
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}}
 
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</ref> in 128 [[Family (biology)|families]] and 47 [[Superfamily|superfamilies]]. The name is derived from [[Ancient Greek]] [[wikt:λεπίδος|λεπίδος]] (scale) and [[wikt:πτερόν|πτερόν]] (wing). Estimates of species suggest that the order may have more species and may be among the largest two or three orders, along with the [[hymenoptera]] and the [[coleoptera]].<ref name=insencyc>Powell, Jerry A. Lepidoptera (pp. 631-664) in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of  Insects. Academic Press.</ref>
 
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==Characteristics==
 
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Lepidopterans  like all [[holometabola]], undergo [[metamorphosis (biology)|complete metamorphosis]], going through a four-stage [[Biological life cycle|life cycle]] of egg - [[larva]]/[[caterpillar]] - [[pupa]]/[[Pupa#Chrysalis|chrysalis]] - [[imago]]/adult.<ref name=insencyc/> Their lifecycle can include an inactive period or [[diapause]] in any of the pre-adult stages that helps overcome unsuitable environmental conditions.<ref name=insencyc/>
 
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The larvae, [[caterpillar]]s, have a toughened (sclerotized) head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections, 3 pairs of true legs, and additional prolegs (up to 5 pairs). They can be confused with the larvae of [[Symphyta|sawflies]]. Lepidopteran larvae can be differentiated by the presence of crochets on the prolegs which are absent in the [[Symphyta]] (sawflies). Most caterpillars are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.<ref name="dugdale">Dugdale, JS, 1996. Natural history and identification of litter-feeding Lepidoptera larvae (Insecta) in beech forests, Orongorongo Valley, New Zealand, with especial reference to the diet of mice (''Mus musculus''). Journal of The Royal Society of New Zealand, Volume 26, Number 2, pp 251-274</ref>
 
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Adults have two pairs of membranous wings covered, usually completely, by minute [[Scale (zoology)|scales]]. In some species, wings are reduced or absent (often in the female but not the male). Antennae are prominent. In moths, males frequently have more feathery antennae than females, for detecting the female [[pheromone]]s at a distance. The [[Trichoptera]] (caddisflies) which are a sister group of the Lepidoptera have scales, but also possess caudal cerci on the abdomen, a feature absent in the Lepidoptera.<ref name=insencyc/>
 
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Adult [[mouth parts]] prominently include the proboscis formed from maxillary galeae and are adapted for sucking nectar. Some species have reduced mouth parts (some species do not feed as adults), and others have them modified to pierce and suck blood or fruit juices (some Noctuids).<ref name="scoble">Scoble, MJ 1995. The Lepidoptera: form, function and diversity. Oxford, UK: The Oxford University Press; 404 p.</ref> Mandibles are absent in all except the [[Zeugaloptera]] which have chewing mouthparts.<ref name=btj>Borror, D.J., Triplehorn, C.A. Johnson. ( 1989) Introduction to the Study of Insects. 6th ed. Brooks Cole.</ref> The maxillary palpi are reduced and consist of up to five segments. They are conspicuous in some of the more primitive families and are often folded. The labial palpi are more prominent and upward pointed.<ref name="insencyc"/>(See also: [[difference between a butterfly and a moth]])
 
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The three thoracic segments are fused and consist of non-movable sclerites. The wings arise from the meso- and meta-thoracic segments and are similar in size in the primitive groups. In the more recent groups, the meso-thoracic wings are larger with more powerful musculature at their bases and more rigid vein structures on the costal edge. In the Noctuoidea, the metathorax is modified with a pair of tympanal organs. There are a variety of wing coupling mechanisms that connect the forewings and the hindwings. The more primitive groups have an enlarged lobe, jugum, at the base of the forewing that folds under the hindwing in flight. Other groups have a frenulum on the hindwing that hooks under a retinaculum on the forewing. In some groups such as the Psychidae, Lymantriidae, the females are flightless and have reduced wings.<ref name=insencyc/>
 
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The abdominal segments 7-10 or 8-10 are modified to form the external genitalia. The abdomen is connected to the thorax in the more recent families by muscles connectd to projections from the abdominal sternite 2. Paired hearing organs at the base of the abdomen occur in the Pyraloidea and Geometroidea. Males have glandular organs such as expandable hair brushes or tufts, or as thin-walled, eversible sacs (coremata), from the intersegmental membranes. The genitalia are complex and provide the basis for species discrimination in most families and also in family identification.<ref name=insencyc/>
 
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Primitive groups have a single genital aperture near the end of the abdomen through which both copulation and egg laying occur. This character is used to designate the Monotrysia. Hepialidae and related families have an external groove that carries sperm from the copulatory opening (gonopore) to the (ovipore) and are termed exoporian. The remaining groups have an internal duct that carry sperm and form the Ditrysia, with two distinct openings each for copulation and egg-laying.<ref name=insencyc/>It also has antennaes.
 
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== Families ==
 
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[[Image:Lepidoptere(s).jpg|thumb|left|140px|[[Forester Moth]] (Zygaenidae)]]
 
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There are about 130 families in this order with variations depending on the taxonomic treatment (see the family template box at the bottom of this section).
 
  
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The Lepidoptera are divided into several suborders, the largest being [[Glossata]], the vast majority of which are [[Ditrysia]].
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The larvae, caterpillars, have a toughened head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections, 3 pairs of true legs, and additional prolegs (up to 5 pairs).  
  
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Several other classifications of lepidopteran families are used in older literature. These include the [[Rhopalocera]] (club-horned) consisting of what are commonly called butterflies and the [[Heterocera]] (varied-horned) consisting of the moths. However, Rhophalocera is a natural ([[monophyletic]]) group, while Heterocera is a [[paraphyletic]] assemblage.
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Adults have two pairs of membranous wings covered, usually completely, by minute scales. In some species, wings are reduced or absent (often in the female but not the male). Antennae are prominent. In moths, males frequently have more feathery antennae than females, for detecting the female pheromones at a distance.  
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Another non-standard classification separates the Lepidoptera into [[Microlepidoptera]] for the smaller species (mostly moths) and [[Macrolepidoptera]] for the larger species.
 
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==Evolution==
 
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===History of study===
 
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Linnaeus in ''Systema Naturae'' (1758) recognized three divisions of the Lepidoptera: ''Papilio, Sphinx,'' and ''[[Phalaena]]'' with seven subgroups in ''Phalaena''.<ref name="scoble"/> These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Denis & [[Ignaz Schiffermüller]] (1775), Fabricius (1775) and [[Pierre André Latreille]] (1796). [[Jacob Hübner]] described many genera, and the Lepidopteran genera were catalogued by [[Ferdinand Ochsenheimer|Ochsenheimer]] and [[Georg Friedrich Treitschke|Treitschke]] in a series of volumes on the Lepidopteran fauna of Europe published between 1807 and 1835.<ref name="scoble"/> G.A.W. Herrich-Schaffer (several volumes, 1843-1856), and [[Edward Meyrick]] (1895) based their classifications primarily on wing venation. Sir [[George Francis Hampson]] worked on the 'microlepidoptera' during this period.
 
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Among the first entomologists to study fossil insects and their evolution was Samuel H Scudder (1837-1911), who worked on butterflies.<ref name="grimaldi">Grimaldi, D, and M S Engel, 2005. Evolution of the Insects. Cambridge University Press.</ref> He published a study of the Florissant deposits of Colorado. Andreas V. Martynov (1879-1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.<ref name="grimaldi"/>
 
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Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.<ref name="scoble"/> [[Willi Hennig]] (1913-1976) developed the [[cladistic]] methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis  studied the relationships among [[monotrysia]]n families  and Kristensen worked more generally on insect [[phylogeny]] and higher Lepidoptera too.<ref name="scoble"/><ref name="grimaldi"/>. While it is often found that DNA-based phylogenies  differ from those based on [[Morphology (biology)|morphology]], this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.<ref name="grimaldi"/>
 
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Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not [[monophyly|monophyletic]]: Microlepidotera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.<ref name="scoble"/>
 
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===Fossils===
 
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Lepidoptera tend not to be as common as some other insects in the habitats that are most conducive to fossilization, such as lakes and ponds, and their juvenile stage  has only the head capsule as a hard part that might be preserved. Yet there are fossils, some preserved in amber and some in very fine sediments. [[Leaf miner|Leaf mines]] are also seen in fossil leaves, although the interpretation of them is tricky.<ref name="grimaldi"/>
 
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The earliest fossil is ''Archaeolepis mane'' from the Jurassic, about 190 million years ago in Dorset, UK.<ref name="grimaldi"/> It consists of wings and shows scales with parallel grooves under a scanning electron microscope and the characteristic wing venation pattern shared with [[Trichoptera]].<ref name="grimaldi"/> Only 2 more sets of Jurassic Lepidopteran fossils have been found, and 13 sets in the Cretaceous.<ref name="grimaldi"/> From there, many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber.
 
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===Phylogeny===
 
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{{userboxtop|toptext=&nbsp;}}
 
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{{Clade|style=font-size:75%;line-height:75%
 
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|1={{clade
 
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|label1=Apoditrysia
 
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|1={{clade
 
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|Label1=Obtectomera
 
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|1={{clade
 
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|label1=Macrolepidoptera
 
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|1= {{clade
 
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|label1=Rhopalocera
 
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|1= {{clade
 
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|1=[[Papilionoidea]]&nbsp;(true&nbsp;butterflies)
 
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|2=[[Hesperiidae]]&nbsp;(skippers)
 
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|3=[[Hedylidae]]&nbsp;(American&nbsp;moth-butterflies)
 
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}}
 
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|2= {{clade
 
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|1=[[Geometroidea]]&nbsp;(geometer&nbsp;moths)
 
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|2=[[Drepanoidea]]&nbsp;(hooktip&nbsp;moths)
 
 
}}
 
}}
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|3=[[Cimeliidae]]&nbsp;(gold&nbsp;moths)
 
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|4=[[Callidulidae]]&nbsp;(old&nbsp;world&nbsp;butterfly&nbsp;moths)
 
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|5=[[Noctuoidea]]&nbsp;(Owlet,&nbsp;tiger&nbsp;moths)
 
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|6= {{clade
 
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|1=[[Bombycoidea]]&nbsp;(Silk&nbsp;moths,&nbsp;hawk&nbsp;moths)
 
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|2=[[Lasiocampoidea]]&nbsp;(lappet&nbsp;moths)
 
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|3=[[Mimallonidae]]&nbsp;(sackbearer&nbsp;moths)
 
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}}
 
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}}
 
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|2=[[Thyrididae]]  (picture&nbsp;winged&nbsp;leaf&nbsp;moths)
 
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|3=[[Hyblaeidae]]&nbsp;(teak&nbsp;moths)
 
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|4=[[Copromorphoidea]]&nbsp;(fruitworm&nbsp;moths)
 
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|5=[[Pyraloidea]]&nbsp;(snout&nbsp;moths)
 
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|6=[[Immidae]]
 
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|7=[[Whalleyana]]
 
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}}
 
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|2=[[Tortricoidea]]&nbsp;(leafrollers)
 
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|3=[[Zygaenoidea]]&nbsp;(burnet&nbsp;moths)
 
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|4=[[Pterophoridae]]&nbsp;(plume&nbsp;moths)
 
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|5=[[Alucitoidea]]&nbsp;(many-plumed&nbsp;moths)
 
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|6=[[Epermeniidae]]&nbsp;(fringe-tufted&nbsp;moths)
 
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|7=[[Schreckensteinia]]&nbsp;(bristle legged&nbsp;moths)
 
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|8=[[Choreutidae]]&nbsp;(metalmark&nbsp;moths)
 
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|9=[[Urodidae]]&nbsp;(false-burnet&nbsp;moths)
 
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|10={{clade
 
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|1=[[Sesioidea]]&nbsp;(clearwing&nbsp;moths)
 
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|2=[[Cossoidea]]&nbsp;(carpenter&nbsp;moths)
 
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}}
 
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}}
 
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|2=[[Gelechioidea]]&nbsp;(twirler&nbsp;moths)
 
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|3=[[Yponomeutoidea]]&nbsp;(ermine&nbsp;moths)
 
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|4=[[Gracillarioidea]]&nbsp;(leafminers)
 
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|5=[[Tineoidea]]&nbsp;(bagworm&nbsp;moths)
 
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}}
 
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}}
 
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<center><small>A proposed phylogeny of the principal lepidopteran groups.<ref>[http://tolweb.org/Ditrysia/11868 Tree of Life] Accessed January 2007</ref></small></center>
 
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{{userboxbottom}}
 
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It has long been noted that the Lepidoptera and the [[Trichoptera]] (caddisflies) share many similarities that are lacking in other insect orders. Among these are:
 
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* females, rather than males, are [[ZW sex-determination system|heterogametic]] (i.e. their sex chromosomes differ)
 
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* dense [[setae]] on the wings (modified into scales in Lepidoptera)
 
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* a particular wing venation pattern on the forewings
 
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* larvae with mouth structures and glands to make and manipulate silk.<ref name="grimaldi"/>
 
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Thus the two sister orders are grouped into the [[Amphiesmenoptera]]. The group probably evolved in the [[Jurassic]], diverging  from the extinct Necrotaulidae.<ref name="grimaldi"/> Lepidoptera differ from the [[Trichoptera]] in several features, including wing venation, form of the scales on the wings, loss of the [[cerci]], loss of an [[ocellus]], and changes to the legs.<ref name="grimaldi"/>
 
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The oldest, most ''basal'' lineages of Lepidoptera have as adults, not the curled tongue or [[proboscis]] characteristic of most members of the order, but chewing [[mandibles]] ([[Micropterigidae]], [[Agathiphagidae]] and [[Heterobathmiidae]]). Micropterigidae larvae feed on decaying leaves (much like the [[Trichoptera]]), [[fungi]], [[liverwort]]s or live leaves.<ref name="scoble"/> The adults chew pollen or spores of ferns. In the Agathiphagidae, larvae feed inside  seeds of [[kauri pine]]s, and in Heterobathmiidae the larvae mine leaves of ''[[Nothofagus]]'', the southern beech. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon just before adult emergence.<ref name="scoble"/>
 
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The [[Eriocraniidae]] have a short coiled proboscis in the adult stage, and retain mandibles for the purpose of escaping the cocoon, but they are non-functional thereafter.<ref name="scoble"/> They, and most of the other non-ditrysian families, are primarily [[leaf miner]]s in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.<ref name="grimaldi"/>
 
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With the evolution of the [[Ditrysia]] in the mid-[[Cretaceous]], there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one [[cloaca]], or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, [[Gracillarioidea]] and [[Gelechioidea]] are mostly leaf miners, but more recent lineages feed externally. In the [[Tineoidea]], most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.<ref name="grimaldi"/>
 
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The [[Yponomeutoidea]] is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.<ref name="grimaldi"/> They evolved about the time that flowering plants underwent an expansive [[adaptive radiation]] in the mid-[[Cretaceous]], and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved [[co-evolution]] or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
 
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In the so-called "[[macrolepidoptera]]", which constitutes about 60% of Lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.<ref name="grimaldi"/> Many also have [[tympanal organ]]s, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.<ref name="grimaldi"/>
 
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The main lineages in the macrolepidoptera are the [[Noctuoidea]], [[Bombycoidea]], [[Lasiocampidae]], [[Mimallonoidea]], [[Geometroidea]] and [[Rhopalocera]]. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a [[monophyletic]] group.<ref name="grimaldi"/> The Rhopalocera, comprising the [[Papilionoidea]] (Butterflies), [[Hesperioidea]] (skippers), and the [[Hedyloidea]] (moth-butterflies), are the most recently evolved.<ref name="scoble"/> There is quite a good fossil record for this group, with the oldest skipper about 56 million years old.<ref name="grimaldi"/>
 
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== External links ==
 
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{{Wikispecies|Lepidoptera}}
 
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{{Wikibookspar|Dichotomous Key|Lepidoptera}}
 
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{{commonscat|Lepidoptera}}
 
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*[http://www.lepidoptera.forumactif.com/ Forum Lepidoptera]
 
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*[http://www.lepidoptera.pl European Butterflies and Moths]
 
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*[http://www.lepidopterology.com/index.htm Lepidopterology.com]
 
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*[http://www.leps.it/ Moths and Butterflies of Europe and North Africa]
 
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*[http://www.ukmoths.force9.co.uk/  British Butterflies and Moths]
 
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*[http://home.hccnet.nl/bernard.fransen/0testsite/0testsite/ Photography of European Butterflies and Moths]
 
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*[http://www.leps.nl/ Butterflies and Moths in the Netherlands]
 
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*[http://www.cirrusimage.com/butterflies_of_North_America.htm Lepidoptera of North America]
 
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*[http://www.goliathus.cz/en/museum-lepidoptera-6.html Lepidoptera] in [http://www.goliathus.cz/en/museum-homepage-0.html online insect museum]
 
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*[http://www.imagenesdeasturias.com/web/panoramicas_graonet/mariposas_de_asturias/index_mariposas.htm Butterflies of Asturias - Spain ]
 
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*[http://www.inra.fr/internet/Produits/PAPILLON/index.htm Lepidoptera of French Antilles]
 
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*[http://pisum.bionet.nsc.ru/kosterin/korgor/index.htm Butterflies of Asian Russia]
 
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*[http://yutaka.it-n.jp/papi.html Butterflies from Indo_China]
 
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*[http://www.mbarnes.force9.co.uk/jamaicamoths/jamaicahome.htm Moths of Jamaica]
 
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*[http://bcrc.bio.umass.edu/kunkel/Moths/ Historic Moth illustrations]
 
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*{{ITIS|ID=117232|taxon=Lepidoptera}}
 
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==Cited references==
 
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<references/>
 
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==Other references==
 
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*Kristensen, NP (Ed.). 1999. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. ''Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches / Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom.'' Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
 
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*Nye, IWB & DS Fletcher, 1991. ''Generic Names of Moths of the World.'' Volume 6: xxix + 368 pp. Trustees of the British Museum (Natural History), London.
 
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*''Firefly Encyclopedia of Insects and Spiders'', edited by Christopher O'Toole, ISBN 1-55297-612-2, 2002
 
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* F. Nemos: ''Europas bekannteste Schmetterlinge. Beschreibung der wichtigsten Arten und Anleitung zur Kenntnis und zum Sammeln der Schmetterlinge und Raupen.'' Oestergaard Verlag, Berlin, ca. 1895, http://hdl.handle.net/10013/epic.28790.d001 (pdf, 77 MB).
 
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{{Lepidoptera}}
 
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[[Category:Insects]]
 
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[[Category:Lepidoptera| ]]
 
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{{link FA|sl}}
 
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[[ar:قشريات الجناح]]
 
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[[id:Kupu-kupu dan ngengat]]
 
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[[ca:Lepidòpter]]
 
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[[eo:Papilio]]
 
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[[fa:پولک‌بالان]]
 
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[[fr:Lepidoptera]]
 
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[[fy:Flinters]]
 
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[[ko:나비목]]
 
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[[hr:Leptiri]]
 
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[[he:פרפראים]]
 
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[[ka:ქერცლფრთიანები]]
 
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[[ht:Papiyon]]
 
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[[lv:Tauriņu kārta]]
 
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[[lt:Drugiai]]
 
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[[simple:Lepidoptera]]
 
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Revision as of 01:37, 12 March 2008

Lepidoptera

Lepidoptera

Butterflies, Moths and Skippers (Lepidoptera)

Order: Lepidoptera

Description: Lepidopterans undergo complete metamorphosis, going through a four-stage life cycle:

  1. egg
  2. larva/caterpillar
  3. pupa/chrysalis
  4. imago/adult.
The larvae, caterpillars, have a toughened head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections, 3 pairs of true legs, and additional prolegs (up to 5 pairs). Adults have two pairs of membranous wings covered, usually completely, by minute scales. In some species, wings are reduced or absent (often in the female but not the male). Antennae are prominent. In moths, males frequently have more feathery antennae than females, for detecting the female pheromones at a distance.

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