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 The Clipper Parthenos sylvia
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See Taxonomy of Lepidoptera and Lepidopteran diversity. A butterfly is an Insect of the order Lepidoptera. Like all Lepidoptera butterflies are notable for their unusual life cycle with a A skipper is a Butterfly of the family Hesperiidae (superfamily Hesperioidea named after their quick darting flight habits A moth is an Insect closely related to the Butterfly, both being of the order Lepidoptera. The Jurassic is a geologic period and system that extends from about Ma (million years ago to Ma that is from the end of the Triassic to the beginning Arthropods are Animals belonging to the Phylum Arthropoda (from Greek ἄρθρον arthron, " Joint " Insects ( Class Insecta) are a major group of Arthropods and the most diverse group of Animals on the Earth with over a million described Pterygota is a subclass of Insects that includes the winged insects Neoptera is a classification group that includes almost all the winged Insects specifically those that can flex their wings over their abdomens The Endopterygota, also known as Holometabola, are Insects of the subclass Pterygota which go through distinctive Larval Pupal Carl Linnaeus (Latinized as Carolus Linnaeus, also known after his ennoblement as, May 23 new style (13 May old style 1707 who laid the foundations for The book Systema Naturae was one of the major works of the Swedish doctor of medicine Carolus Linnaeus. This is a taxonomy of the order Lepidoptera down to family level Suborder Aglossata Superfamily Agathiphagoidea This is a list of the Diversity of the Lepidoptera showing the estimated number of genera and species described for each superfamily and where available family |
Lepidoptera is an order of insect that includes moths and butterflies. This article is about the taxonomic rank for the sequence of species in a taxonomic list see Taxonomic order In scientific classification used Insects ( Class Insecta) are a major group of Arthropods and the most diverse group of Animals on the Earth with over a million described A butterfly is an Insect of the order Lepidoptera. Like all Lepidoptera butterflies are notable for their unusual life cycle with a It is one of the most speciose orders in the class Insecta, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies. A class is the Taxonomic rank in the Biological classification of organisms in Biology below phylum and above order. Insects ( Class Insecta) are a major group of Arthropods and the most diverse group of Animals on the Earth with over a million described A moth is an Insect closely related to the Butterfly, both being of the order Lepidoptera. A butterfly is an Insect of the order Lepidoptera. Like all Lepidoptera butterflies are notable for their unusual life cycle with a A skipper is a Butterfly of the family Hesperiidae (superfamily Hesperioidea named after their quick darting flight habits Hedylidae, the "American moth-butterflies" is a family of Insects in the Lepidopteran order representing the superfamily Members of the order are referred to as lepidopterans. A person who collects or studies this order is referred to as a lepidopterist. A lepidopterist is a person who catches and collects studies or simply observes (see Butterfly watching) lepidopterans, members of an order encompassing This order has more than 180,000 species[1] in 128 families and 47 superfamilies. In Biology, a species is one of the basic units of Biological classification and a Taxonomic rank. In Biological classification, family ( Latin Taxonomic rank ( rank, category, taxonomic category is an abstract term used in the Scientific classification, or Taxonomy, of organisms The name is derived from Ancient Greek λεπίδος (scale) and πτερόν (wing). The Ancient Greek language is the historical stage in the development of the Hellenic language family spanning the Archaic (c 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. Hymenoptera is one of the larger orders of Insects comprising the sawflies, Wasps Bees and Ants The name refers to Beetles are the group of Insects with the largest number of known Species. [2]
Contents |
General Characteristics
Lepidopterans like all holometabola, undergo complete metamorphosis, going through a four-stage life cycle: egg, larva/caterpillar, pupa/chrysalis, and imago/adult. The Endopterygota, also known as Holometabola, are Insects of the subclass Pterygota which go through distinctive Larval Pupal Metamorphosis is a Biological process by which an Animal physically develops after Birth or hatching involving a conspicuous and relatively A life cycle is a period involving 1 Generation of an Organism through means of Reproduction, whether through Asexual reproduction or Sexual A larva ( Latin; plural larvae) is a juvenile form of Animal with indirect development, undergoing Metamorphosis (for example Caterpillars are the Larval form of a member of the order Lepidoptera (the Insect order comprising butterflies and Moths A pupa ( Latin pupa for doll pl pupae or pupas) is the life stage of some Insects undergoing transformation A pupa ( Latin pupa for doll pl pupae or pupas) is the life stage of some Insects undergoing transformation In Biology, the imago is the last stage of development of an Insect, after the last Ecdysis of an incomplete metamorphosis, or after emergence [2] Their lifecycle can include an inactive period or diapause in any of the pre-adult stages that helps them to overcome unsuitable environmental conditions. Diapause is a Physiological state of Dormancy with very specific triggering and releasing conditions [2]
The larvae, caterpillars, 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). Caterpillars are the Larval form of a member of the order Lepidoptera (the Insect order comprising butterflies and Moths They can be confused with the larvae of sawflies. Symphyta redirects here For the Moth Genus, see Symphyta (genus. Lepidopteran larvae can be differentiated by the presence of crochets on the prolegs which are absent in the Symphyta (sawflies). Symphyta redirects here For the Moth Genus, see Symphyta (genus. Most caterpillars are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores. [3]
Adults have two pairs of membranous wings covered, usually completely, by minute scales. In most biological nomenclature a scale ( Greek lepid, Latin squama) is a small rigid plate that grows out of an Animal 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. A pheromone (from Greek φέρω phero "to bear" + ‘ορμόνη " Hormone " is a Chemical that triggers a natural 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. Caddisflies, sedge-flies or rail flies (Order Trichoptera, from Greek trich, "hair" and ptera, "wings" [2]
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). [4] Mandibles are absent in all except the Zeugaloptera which have chewing mouthparts. [5] 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. [2](See also: difference between a butterfly and a moth)
The three thoracic segments are fused and consist of non-movable sclerites. A common classification of the Lepidoptera involves their differentiation into butterflies and Moths Butterflies are a natural Monophyletic group often 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. [2]
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. [2]
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. [2]It also has antennaes.
Families
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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). The Zygaenidae are a family of Lepidoptera ( Moths typically day-flying with a slow fluttering flight and with rather clubbed antennae They generally
The Lepidoptera are divided into several suborders, the largest being Glossata, the vast majority of which are Ditrysia. Glossata is the Suborder of the Insect order Lepidoptera that includes all the Superfamilies of moths and butterflies that have a coilable The Ditrysia are a natural group or clade of Insects in the Lepidopteran order containing both butterflies and Moths
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. A butterfly is an Insect of the order Lepidoptera. Like all Lepidoptera butterflies are notable for their unusual life cycle with a A common classification of the Lepidoptera involves their differentiation into butterflies and Moths Butterflies are a natural Monophyletic group often However, Rhophalocera is a natural (monophyletic) group, while Heterocera is a paraphyletic assemblage. A clade is a taxonomic group comprising a single Common ancestor and all the descendants of that ancestor In Phylogenetics, a group of organisms is said to be paraphyletic if the group contains its most recent common ancestor but does not contain all
Another non-standard classification separates the Lepidoptera into Microlepidoptera for the smaller species (mostly moths) and Macrolepidoptera for the larger species. Microlepidoptera is an artificial (ie unranked and not Monophyletic) grouping of Moth families commonly known as the 'smaller moths' ( Micro, Lepidoptera Macrolepidoptera is a group within the Insect order Lepidoptera.
Evolution
Butterflies have evolved over time by the myriad amounts of species in the order. Most species are inter-related due to the evolution of the family for millions of years.
History of study
Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx, and Phalaena with seven subgroups in Phalaena. The book Systema Naturae was one of the major works of the Swedish doctor of medicine Carolus Linnaeus. Phalaena was originally a subdivision of Lepidoptera, created by Carolus Linnaeus, and included Moths in general [4] 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). Johann Nepomuk Cosmas Michael Denis ( 27 September 1729 - 29 September 1800) was an Austrian Poet, Bibliographer Ignaz Schiffermüller (1727 - 1806 was an Austrian naturalist mainly interested in Lepidoptera. Johan Christian Fabricius ( January 7, 1745 - March 3, 1808) was a Danish Entomologist and economist Pierre André Latreille ( November 20, 1762 - February 6, 1833) was a French entomologist. Jacob Hübner described many genera, and the Lepidopteran genera were catalogued by Ochsenheimer and Treitschke in a series of volumes on the Lepidopteran fauna of Europe published between 1807 and 1835. Jacob Hübner ( June 20, 1761 - September 13, 1826, Augsburg) was a German Entomologist. Ferdinand Ochsenheimer ( 17 March 1767 &ndash 2 November 1822) was a German actor and Entomologist ( Lepidopterist Georg Friedrich Treitschke ( August 29, 1776, Leipzig, Germany &ndash June 4 1842, Vienna, Austria [4] G.A.W. Herrich-Schaffer (several volumes, 1843-1856), and Edward Meyrick (1895) based their classifications primarily on wing venation. Dr Gottlieb August Wilhelm Herrich-Schäffer ( December 17 1799 - April 14 1874) was a German Entomologist. Edward Meyrick ( November 24, 1854 - March 31, 1938) was an English schoolmaster and amateur Entomologist. Sir George Francis Hampson worked on the 'microlepidoptera' during this period and Philipp Christoph Zeller published The Natural History of the Tineinae13 volumes also on 'microlepidoptera'(1855). Sir George Francis Hampson 10th Baronet ( January 14 1860 - October 15 1936) was a British Entomologist. Philipp Christoph Zeller ( April 8, 1808 - March 27, 1883) was a German Entomologist.
Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder (1837-1911), who worked on butterflies. Samuel Hubbard Scudder was an American Entomologist and Palaeontologist. [6] 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. Andrey V Martynov (1879 &ndash 1938 was a Russian entomologist and palaeontologist, a founder of the Russian palaeoentomological school [6] 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. [4] Willi Hennig (1913-1976) developed the cladistic methodology and applied it to insect phylogeny. Emil Hans Willi Hennig ( April 20, 1913 in Dürrhennersdorf/ Oberlausitz – November 5, 1976 in Ludwigsburg) was a Cladistics is the hierarchical classification of Species based on evolutionary ancestry Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysian families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too. The Monotrysia is a group of Insects in the Lepidopteran order which is not currently considered to be a natural group or clade. [4][6]. While it is often found that DNA-based phylogenies differ from those based on morphology, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies. The term morphology in Biology refers to the outward appearance ( Shape, Structure, Colour, Pattern) of an Organism [6]
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 monophyletic: Microlepidotera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia. A clade is a taxonomic group comprising a single Common ancestor and all the descendants of that ancestor [4]
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 mines are also seen in fossil leaves, although the interpretation of them is tricky. Leaf miner is a term used to describe the Larvae of many different species of Insect which live in and eat the Leaf tissue of plants [6] The earliest fossil is Archaeolepis mane from the Jurassic, about 190 million years ago in Dorset, UK. [6] It consists of wings and shows scales with parallel grooves under a scanning electron microscope and the characteristic wing venation pattern shared with Trichoptera. Caddisflies, sedge-flies or rail flies (Order Trichoptera, from Greek trich, "hair" and ptera, "wings" [6] Only 2 more sets of Jurassic Lepidopteran fossils have been found, and 13 sets in the Cretaceous. [6] From there, many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber.
Phylogeny
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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. Caddisflies, sedge-flies or rail flies (Order Trichoptera, from Greek trich, "hair" and ptera, "wings" Among these are:
- females, rather than males, are heterogametic (i. The ZW sex-determination system is a system that determines the sex of offspring in Birds, some Fish, and some Insects (including Butterflies e. their sex chromosomes differ)
- dense setae on the wings (modified into scales in Lepidoptera)
- a particular wing venation pattern on the forewings
- larvae with mouth structures and glands to make and manipulate silk. Seta (plural setae) is a biological term derived from the Latin word for " Bristle " [6]
Thus the two sister orders are grouped into the Amphiesmenoptera. Amphiesmenoptera is an Insect Superorder, established by Willi Hennig in his revision of insect taxonomy for two sister orders Lepidoptera The group probably evolved in the Jurassic, diverging from the extinct Necrotaulidae. The Jurassic is a geologic period and system that extends from about Ma (million years ago to Ma that is from the end of the Triassic to the beginning [6] 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. Caddisflies, sedge-flies or rail flies (Order Trichoptera, from Greek trich, "hair" and ptera, "wings" Cerci (singular cercus) are paired Appendages on the rear-most segments of many Arthropods including Insects and Arachnids but not An ocellus (plural ocelli) is a type of Photoreceptor organ in animals [6]
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). In general a proboscis (from Greek προ pro "before" and βοσκειν boskein "to feed" also known as probiscus The mandible (from Latin mandibula, "jawbone" or inferior maxillary bone forms the lower Jaw and holds the lower teeth in place Micropterigoidea is the superfamily of "mandibulate archaic moths" all placed in the single family Micropterigidae, containing currently 12 living genera Agathiphaga is a Genus of Insects in the order Lepidoptera. This Caddis fly -like lineage of primitive "Kauri moths" The Heterobathmioidea are a superfamily of the Lepidoptera, containing a single genus Heterobathmia. Micropterigidae larvae feed on decaying leaves (much like the Trichoptera), fungi, liverworts or live leaves. Caddisflies, sedge-flies or rail flies (Order Trichoptera, from Greek trich, "hair" and ptera, "wings" A fungus (ˈfʌŋgəs is a eukaryotic Organism that is a member of the kingdom Fungi (ˈfʌndʒaɪ [4] The adults chew pollen or spores of ferns. In the Agathiphagidae, larvae feed inside seeds of kauri pines, and in Heterobathmiidae the larvae mine leaves of Nothofagus, the southern beech. The genus Agathis, commonly known as kauri or dammar, is a relatively small Genus of 21 species of Evergreen Trees in the Nothofagus, also known as the southern beeches, is a genus of about 35 Species of Trees and shrubs native to the temperate oceanic These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon just before adult emergence. [4]
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. A superfamily of insects in the Lepidoptera order restricted to the Holarctic region with six genera (see Davis 1978 Mizukawa et al [4] They, and most of the other non-ditrysian families, are primarily leaf miners in the larval stage. Leaf miner is a term used to describe the Larvae of many different species of Insect which live in and eat the Leaf tissue of plants In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales. [6]
With the evolution of the Ditrysia in the mid-Cretaceous, there was a major reproductive change. The Ditrysia are a natural group or clade of Insects in the Lepidopteran order containing both butterflies and Moths The Cretaceous (kriːˈteɪʃəs, usually abbreviated 'K' for its German translation "Kreide" is a geologic period and system, reaching from the end of 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. In zoological anatomy a cloaca is the posterior opening that serves as the only such opening for the intestinal and urinary tracts of certain animal species ) Of the early lineages of Ditrysia, Gracillarioidea and Gelechioidea are mostly leaf miners, but more recent lineages feed externally. Gracillarioidea is a large superfamily containing four families of Insects in the order Lepidoptera. Gelechioidea is the superfamily of moths that includes case bearers and relatives In the Tineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage. Tineoidea is the superfamily of Moths that includes clothes moths bagworms and relatives [6]
The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants. Yponomeutoidea is the Superfamily of Ermine Moths and Relatives Familiae Acrolepiidae Bedelliidae [6] 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. An adaptive radiation is a rapid Evolutionary radiation characterized by an increase in the morphological and ecological diversity of a single rapidly diversifying lineage The Cretaceous (kriːˈteɪʃəs, usually abbreviated 'K' for its German translation "Kreide" is a geologic period and system, reaching from the end of Whether the processes involved co-evolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together. In a broad sense biological co-evolution is "the change of a biological object triggered by the change of a related object"
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. Macrolepidoptera is a group within the Insect order Lepidoptera. [6] Many also have tympanal organs, that allow them to hear. A Tympanal organ is a hearing organ in Insects, consisting of a membrane ( tympanum) stretched across a frame backed by an air sac These organs evolved eight times, at least, because they occur on different body parts and have structural differences. [6] The main lineages in the macrolepidoptera are the Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea and Rhopalocera. Noctuoidea is the Superfamily of noctuid ( Latin "night owl" or "owlet" Moths and has the largest number of Species Bombycoidea is a Superfamily of Moths. It contains the silk moths emperor moths sphinx moths and relatives The Lasiocampidae family of Moths are also known as eggars, snout moths or lappet moths. Mimallonoidea is the superfamily of sack bearer moths containing the single family Mimallonidae. Geometroidea is the superfamily of geometrid moths in the Lepidoptera. A butterfly is an Insect of the order Lepidoptera. Like all Lepidoptera butterflies are notable for their unusual life cycle with a Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group. A clade is a taxonomic group comprising a single Common ancestor and all the descendants of that ancestor [6] The Rhopalocera, comprising the Papilionoidea (Butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved. The Superfamily Papilionoidea (from the word "papilion" meaning "butterfly" contains all the butterflies except for the skippers which A skipper is a Butterfly of the family Hesperiidae (superfamily Hesperioidea named after their quick darting flight habits Hedylidae, the "American moth-butterflies" is a family of Insects in the Lepidopteran order representing the superfamily [4] There is quite a good fossil record for this group, with the oldest skipper about 56 million years old. [6]
Cited references
- ^ The Lepidoptera Taxome Project Draft Proposals and Information. Centre for Ecology and Evolution, University College London. Retrieved on 2007-03-05. Year 2007 ( MMVII) was a Common year starting on Monday of the Gregorian calendar in the 21st century. Events 363 - Roman Emperor Julian moves from Antioch with an army of 90000 to attack the Sassanid Empire, in a
- ^ a b c d e f g h Powell, Jerry A. Lepidoptera (pp. 631-664) in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of Insects. Academic Press.
- ^ 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
- ^ a b c d e f g h i j Scoble, MJ 1995. The Lepidoptera: form, function and diversity. Oxford, UK: The Oxford University Press; 404 p.
- ^ Borror, D. J. , Triplehorn, C. A. Johnson. ( 1989) Introduction to the Study of Insects. 6th ed. Brooks Cole.
- ^ a b c d e f g h i j k l m n o p q r Grimaldi, D, and M S Engel, 2005. Evolution of the Insects. Cambridge University Press.
- ^ Tree of Life Accessed January 2007
Other references
- 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.
- 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.
- Firefly Encyclopedia of Insects and Spiders, edited by Christopher O'Toole, ISBN 1-55297-612-2, 2002
- 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).
External links
- Forum Lepidoptera
- European Butterflies and Moths
- Lepidopterology.com
- Moths and Butterflies of Europe and North Africa
- British Butterflies and Moths
- Butterflies of Bulgaria
- Photography of European Butterflies and Moths
- Butterflies and Moths in the Netherlands
- Lepidoptera of North America
- Lepidoptera in online insect museum
- Butterflies of Asturias - Spain
- Lepidoptera of French Antilles
- Butterflies of Asian Russia
- Butterflies from Indo_China
- Moths of Jamaica
- Moths of Japan Excellent photographs. In Japanese but uses binomial names
- Historic Moth illustrations
- Lepidoptera (TSN 117232). Integrated Taxonomic Information System. Integrated Taxonomic Information System ( ITIS) is a partnership designed to provide consistent and reliable information on the Taxonomy of biological Species
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