taxonID	type	description	language	source
039C87C3FFCFDC20FF3EE367FC0EFAD2.taxon	description	A detailed reconstruction of the lepidopteran ground plan was presented by Kristensen (1984) and updated in Kristensen & Skalski 1998). Several apomorphies arguably unique to the Lepidoptera were identified, and another set of groundplan traits was identified as probable synapomorphies with the Trichoptera, i. e., autapomorphies of the Amphiesmenoptera. The great majority of the characters in question occurs in the adult insect. Subsequent work has suggested little in the way of change. The following morphological character states are currently considered likely lepidopteran groundplan autapomorphies: Adult (numbers in brackets refer to figured details in Fig. 1): Median ocellus lost (1). Corporotentorium with posteromedian process, accommodating insertions of ventral neck muscles (2). Intercalary sclerite present laterally in membrane between antennal scapus and pedicellus (3). Maxillary palp with points of flexion between segments 1 / 2 and 3 / 4; segment 4 longest; intrinsic palp musculature not comprising antagonistic pairs (4). Craniostipital muscle present, slender and running close to craniocardinal muscle (5). Postlabium an arched sclerite with long piliform scales (6). Terminal segment of labial palp with group of sensilla in depression (' vom Rath's organ') (7). Labral nerve and frontal ganglion connective separating immediately at their origin on the tritocerebrum (9). Nervus recurrens running inside aorta until reaching retrocerebral complex (10). Laterocervical sclerite with proprioceptive ' hair plate' close to anterior apex (11). Prothoracic endoskeleton with prominent free arm arising from bridge between sternum and lower posterior corner of pleuron (12). Mesothorax with ' tergopleural apodeme' issued from upper part of pleural suture and accommodating insertion of a tergopleural muscle (13). Metathorax with ' prescutal arm' (14). Fore tibia with movable ' epiphysis' on inner surface, and with at most a single spur (15). Wings with dense covering of broad scales (16). Metathoracic spiracle with single, anteriorly situated, external lip (17). Tergum I extensively desclerotized, with concomitant loss of external layer of ' short' dorsolongitudinal I / II muscles (18). Tergum I with lateral lobes extending posteroventral to articulate with anterior corners of sternum II (19). Male ' valve' (gonopod) primarily undivided (20). Phallic protractor muscle originating inside gonopod (‘ valve’) (21). Cerci lacking in both sexes (22). Abdominal nerve cord with at most five ganglionic masses, and unpaired connectives (23). Apyrene sperm present (24). Larva: Pleurostome elongated, craniocardinal articulation far behind mandibular base. Maxillary palp with no more than 3 segments. Absence of a dorsolongitudinal muscle on the adult's salivarium (8) has been considered an additional lepidopteran autapomorphy, but unpublished observations on nannomecopterans (R. Beutel, pers. comm.) and trichopterans render it uncertain that a presence of a formation of this kind is actually the groundplan condition in the Mecopterida. In the light of this robust morphological support for the monophyly of the Lepidoptera, inclusive of the Micropterigoidea, the assignment (Chapman 1917, Hinton 1946, 1958) of the latter to a separate order Zeugloptera is now of historical interest only. Lepidoptera monophyly has also been supported consistently in available molecular analyses with relevant taxon sampling. The following lepidopteran groundplan traits are apparently autapomorphic of the superorder Amphiesmenoptera. Adult (Roman numerals are details in Fig. 1): Prelabium fused with hypopharynx (I). Lower posterior corner of laterocervicale produced towards the prosternum (II). Pronotum with paired setose ' warts' (III). Prothoracic episterna with unique suture pattern (IV). Secondary furcal arms of pterothorax fused with posterior margins of corresponding epimera (V). Metathorax with setose, presumably proprioceptive, sclerite in wing base membrane behind / below subalare (VI). Pretarsus above claw with ' pseudempodium' (strong seta on socket) (VII). Wings with dense vestiture of setae (forerunners of the lepidopteran scales) (VIII). Fore wing anal veins looping up into double-Y formation (IX). One ventral (tentorial) neck muscle originating on fore coxa (X). Conical furcopleural muscle in mesothorax with broad end on pleural ridge (XI). Paired glands opening on sternum V (XII). Male segment IX with tergum and sternum fused into closed ring (XIII). Anterior margin of female segments VIII and IX with long rod-like apodemes accommodating insertions of protractor and retractor muscles of extensible oviscapt (XIV). Note: recent work shows that the interpretation of the female postabdomen in the lowest Amphiesmenoptera is more problematical than hitherto believed, and the apodemes in question may not all be homologous (Kristensen 2003); a particularly intriguing question is whether a three-apophysis-pair configuration (with both dorsal and ventral apophyses originating on VIII) could prove ancestral in Amphiesmenoptera, since three pairs are present in Agathiphaga as well as in the enigmatic recently described caddisfly Fansipangana (Mey 1996). Ventral diaphragm muscles inserting on the nerve cord (XV). Female sex heterogametic (XVI). Chromosome number unusually high (basic number 30 – 31), chromosomes holocentric and oogenesis achiasmatic (XVII). Spermatozoa with outer accessory filaments thickened, filled with proteinaceous and glycogen-like material (XVIII). Larva: stemmata each with one crystalline cone cell transformed into primary pigment cell, hence in transverse section the cone is seen to be only tripartite. Prelabium and hypopharynx fused into composite lobe with silk gland orifice on apex. Evidence suggests that larvae of members of the amphiesmenopteran stem lineage were ' soil animals', living in wet conditions, like those of most extant Lepidoptera-Micropterigidae. The step from here to a truly aquatic lifestyle, which is autapotypic of immature Trichoptera, is but a small one. There is every reason to believe that these larvae were free-living, for the use of silk as part of pre-pupating larval behaviour first evolved, as far we know, in the stem lineage of the Neolepidoptera.	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
039C87C3FFC9DC2DFF3EE4EAFEE5FE42.taxon	description	Linnaeus based his classification on features of the antennae and the wings, particularly the position of the wings at rest, and on whether the insects flew by day or by night. He also took account of the mouthparts. He noted that the antennae of butterflies were clubbed, and that some members of what he called ‘ Bombyces’ lacked a tongue. A considerable variety of features of larvae and adults is to be found in his species descriptions. Lepidopteran classification developed from the Linnaean foundation in several ways. Numerous additional species were described, as discussed elsewhere in this paper and Linnaeus’ divisions and subdivisions were further divided and formal categorical ranks were established for them. Moreover, many supraspecific taxa were added to those introduced by Linnaeus. Primary types of many Linnean Lepidoptera species survive to this day, and thus remain available for study. Linnaeus’ own collection is housed in the Linnean Society of London (see http: // www. linnean. org /); examples are shown in Figs 2 – 3. Prior to Linnaeus’ death, and just under twenty years after the publication of Systema Naturae, Denis & Schiffermüller (1775) produced a work on the Lepidoptera of the Vienna area. Their system was based on Linnaeus’ classification, but they added further observations on structure and examined many additional species. They subdivided the Linnaean groups and based the names of many of the subdivisions on both larva (e. g., Larvae Punctatae) and adult (e. g., Ph [alaenae] Geometrae Unicolores). Building on these foundations, the Danish entomologist Fabricius, who was a student of Linnaeus, described many new species and named some of the Linnaean subgroups (Fabricius 1775 - and several later works). Fabricius’ impact on Lepidoptera taxonomy (indeed insect taxonomy broadly) was notable. Slightly later, Latreille (1796) added further supraspecific groups, most of which represent genera in the modern sense.	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
039C87C3FFC9DC2DFF3EE4EAFEE5FE42.taxon	description	With collections being made from beyond Europe, a further dimension to the classification was added. Apart from the fact that they described numerous new genera and species, Edward Meyrick (particularly for microlepidopterans) and Sir George Hampson (particularly for macrolepidopterans) had a marked impact on the higher classification of the order. Meyrick (1895) proposed that the Lepidoptera should be divided into ten ‘ phyla’. Meyrick, like Herrich-Schäffer, relied to a significant extent on wing venation because he considered the pattern to be of low adaptive (‘ physiological’) value and likely therefore to be little altered by external factors. Hampson’s contribution to Lepidoptera higher classification focused mainly on Noctuoidea (see Kitching 1984 for an analysis), Pyraloidea, and Thyridoidea. Since the mid nineteenth century many other systems have caused the earlier classifications to be modified and expanded. Some writers emphasized other character sets e. g., from larvae (Fracker 1915) or pupae (Mosher 1916). Probably the major contribution to the higher classification of the order concerns the division above the superfamily. Landmark works were written by Börner (1925, 1939) who proposed a fundamental division of the Lepidoptera into Monotrysia and Ditrysia on the basis of the structure of female genitalia. The Monotrysia (sensu Börner), which included only 5 percent of the Lepidoptera, are not monophyletic. But Börner’s recognition of the Ditrysia as a natural group was an important step in understanding the phylogenetic structure of the order. So also was his appreciation of the systematic value of many morphological characters in the classification and diagnosis of lepidopteran families and superfamilies. Building on the works of Börner as well as of Kiriakoff (1948) and in particular Hinton (1946), Hennig (1953) outlined the relationships of the most basal lineages within the order in a benchmark article, which foreshadowed subsequent work on this subject. One hundred and twenty-four lepidopteran families (52 of them with a subfamily classification) were recognized in the recent treatment of the order in the Handbook of Zoology (Kristensen ed. 1998, 2003). These families were grouped into 47 diagnosed superfamilies, and a provisional phylogeny at the superfamily level was presented. About the same time an alternative classification was presented by Heppner (1998), who later (2005) expanded it to tribal level. The two classifications are broadly similar, but some differences exist, partly because Heppner’s classification expressly does not aim to reflect monophyla in the ' Hennigian' (cladistic) sense of phylogenetic systematics. Moreover, the superfamilies and higher-level entities in Heppner’s work were not diagnosed. The superfamily / family / subfamily classification adopted in the Handbook, with some major modifications suggested in subsequent work, is given in Appendix 1. Although the superfamily cladogram printed in the Handbook (and see Fig. 3 in the present paper) has been referred to as representing the “ consensus ” position (Holloway et al. 2001), its provisional nature should be emphasized: its lack of resolution of major branches is conspicuous. It serves, however, as a starting point for some ongoing inquiries into Lepidoptera phylogenetic systematics (see, e. g., the http: // www. tolweb. org / Lepidoptera and http: // www. lepsys. eu / websites). In Fig. 4 the approximate species numbers for superfamilies / superfamily assemblages with more than 1,000 described species is indicated by the width of the clade lines. It is immediately apparent, that the most species-rich lineages are cladistically quite subordinate. This particular phylogenetic pattern was ascribed considerable general significance in Hennig's writings (e. g., 1953, 1966): indeed, the basic diversification pattern within the Lepidoptera remains an unusually instructive example of what has subsequently come to be known as a ' Hennigian comb', with the first diverged extant lineages exhibiting a step-by-step acquisition of the apomorphies which characterize the most subordinate (and successful) groups. Undoubtedly the proportional representation of individual superfamilies will change markedly as descriptive Lepidoptera taxonomy approaches completion. Thus only relatively modest growth is foreseeable in the butterfly and bombycoid assemblages, while particularly strong increases are expected for the Gelechioidea and the Pyraloidea.	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
039C87C3FFC4DC28FF3EE30FFF46F9B2.taxon	description	Gaston (1991 a: 286) was undoubtedly correct when writing that " The Lepidoptera are generally regarded as the best collected and studied of the four major insect orders ". However, on a global scale this general belief is in itself probably incorrect. The advanced levels of butterfly and hawk / emperor moth taxonomy and of inventories of some very restricted Northern Hemisphere Lepidoptera faunas have little bearing on the overall status of systematic lepidopterology. The major tasks for delivering this more comprehensive picture lies in dealing with the world fauna of moths. The sampling of tropical moths has so far been very inadequate, and where samples have been made they have often not been worked up. This situation is significantly explained by the work of Gaston & May (1992). These authors found, on the basis of surveys of insect systematists in North America and the UK, that of the four major insect orders the Lepidoptera actually have the smallest taxonomic workforce. The same situation then pertained also in Australia (pers comm. from the late E. S. Nielsen). This situation is unlikely to have changed today. From their survey, Gaston & May (1992) indicated that about 80 % of present-day insect taxonomists then were based in North America and Europe, a figure unlikely to have changed much subsequently — except, probably, for some E. Asian countries. Taxonomy also has had longstanding strongholds in other industrialized countries such as Japan, Australia and New Zealand. It is therefore to be expected that the faunas will be better known in these regions than elsewhere, but it must also be noted that to several workers based here certain tropical areas (Taiwan, Borneo, Costa Rica, etc.) have recently been of focal interest. Experience shows, that large-scale elucidation of life-histories of the insect fauna of a given area is feasible only for those workers who are at least largely resident in that area. The knowledge of Lepidoptera immatures is, therefore, even more geographically biased than that of the adult insects. ‘ Patriotism’ has also made its mark on our knowledge of distribution in Lepidoptera, particularly in terms of the activities of non-professionals. ' Patriotism' in this context means the sum of linguistic and traditional / cultural barriers, which restrict the activity of a worker to the study of her / his national fauna. Perhaps the most striking evidence of this mechanism is the markedly more detailed knowledge of the moth fauna, particularly of the micro-moths, of Europe compared with that of any N. American area. So many N. American lepidopterists remain ' butterfly workers', since they feel few barriers to extending their interest to the butterfly fauna of the entire Nearctic region. In contrast, the average amateur lepidopterist, at least in some European countries, having exhausted the challenges (at the collectors' level!) presented by the local butterflies, has been more likely to proceed to the study of the macro-moths, and often also the micro-moths, of the national fauna, rather than to butterflies of neighbouring countries. This explains why in some countries even the most minute micro-moths are now known in much greater detail than larger and more striking wasps, flies etc. Having said that, an increasing number of European amateur specialists in recent decades have extended their interests, and descriptive efforts, to Lepidoptera faunas outside their native regions. A provisional survey of the described world fauna of Lepidoptera, by family-group taxa and zoogeographical regions, was published by Heppner (1991) (minor updates are given by Heppner 1998). This survey was based partly on actual counts from the literature, partly on the card indexes in the Natural History Museum (London) and the National Museum of National History (Washington), and partly on estimates (the source is uncertain in many cases). Clearly this task was a difficult one, and Heppner rightly underscored the uncertainties of the figures. Herbulot (1992) and Scoble et al. (1995) highlighted these uncertainties by comparisons with (near-) exact counts of Geometridae, which for some regions and / or subfamilies differed markedly; for instance the figure for Afrotropical species proved to be almost 50 % higher than Heppner's. It so happens, however, that the inexactitudes in Heppner’s figures largely cancel out mutually, hence his grand total for world Geometridae (20,890) is remarkably close to the figure by Scoble et al. (21,144), and similarly his estimated total for world Lepidoptera (146,565) is remarkably close to the estimates given about the same time by Holloway et al. (1987) and Hammond (1992). The number of described species is now probably some 160,000. Table 1 shows a geographical breakdown of Heppner's global numbers of described species. Heppner noted that the Neotropical Lepidoptera fauna is considerably more species-rich than that of the Oriental, and he attempted to find explanations for this observation. He noted, for instance, that the Neotropical region covers by far the larger land area of the two. From Table 1 it is clear that when one combines Heppner's ' Oriental' and ' Australian' regions into an ' Indoaustralian' region (a concept which will be familiar to users of the monumental ' Macrolepidoptera' manuals edited by A. Seitz), the latter has a land area that is very similar to that of the Neotropical region. The two regions are also roughly comparable with respect to north-south extent, and their Lepidoptera diversities, crudely expressed as species per unit area, are then seen to be very similar indeed. The main issue, which arises from the geographical survey is, therefore, the markedly lower species-per-unit-area figure for the Afrotropical region. This pattern, which is found repeatedly in global biodiversity surveys, more than 30 years ago prompted Richards (1973) to refer to Africa as the “ odd man out ”, and it continues to attract biogeographers' attention. To examine patterns of species description in Lepidoptera it is worth considering separately those three major assemblages of the order with which lepidopterists are familiar. Historically, work on these groupings have followed rather different paths. These groupings are 1) The micro-moths, which embrace the more primitive lineages within the order, ' up to', and including, the ' pyraloid grade' superfamilies (mostly small (sometimes minute) moths with larvae usually feeding internally or concealed on their host plants. 2) The macromoths, i. e., the predominantly night-flying members of the Macrolepidoptera, which may, or may not, be monophyletic. The Macrolepidoptera are medium-sized to large insects, the larvae of which are often external feeders. 3) The butterflies, a presumably monophyletic entity that may be cladistically subordinate within the Macrolepidoptera; nearly all adult butterflies are diurnal, and many have particularly colourful wing patterns. Fig. 5 illustrates some trends in the taxonomic research on the three groupings during the latter half of the post-Linnean era, showing the numbers of new species descriptions (A) and new species synonyms (B) published during six three-years periods since the establishment of Zoological Record in 1864. One general trend, which is observable in all curves of new species descriptions, is a marked decline towards the fourth period (mid-twentieth century), followed by a more or less pronounced subsequent rise. However, the concordant curve shapes have different backgrounds, as has already been pointed out by Strong et. al. (1984) in their comments on similar statistics for a variety of phytophagous insect groups. In the case of the butterflies there can be no doubt, that the overall decrease in new species descriptions since a century ago reflects a genuine, even pronounced, drop in the number of species which still remain to be discovered / named. In the cases of the moths the drop reflects the change in the working practice of leading taxonomists, who no longer looked favourably on species descriptions " divorced from revisional or monographic work " (as expressed in Mayr, Linsley & Usinger's influential 1953 manual). Also, examination and specimen preparation procedures had become much more time-consuming, with descriptions / illustrations of genital structures eventually being considered mandatory ingredients in a worthwhile taxonomic study. FIG. URE 5. New species described (upper) and new synonyms (lower) established for micro-moths, macro-moths and butterflies during six 3 - year periods from the start of Zoological Record. The preparation of permanent slide mounts of the genitalia has been a much more widespread practice among Lepidoptera taxonomists than among those working on insects of several other orders (who often are content to observe the organs at dissection microscope magnification only, with preparations stored in glycerol or even mounted dry on cards). This is undoubtedly primarily rooted in tradition, and it may in no small measure be due to the early influence exerted by F. N. Pierce on the British lepidopterists' community, including, eventually, the staff of the Natural History Museum, London. Unquestionably, this practice has enhanced appreciation of small, taxonomically significant structural details. Moreover, carefully prepared permanent mounts often are excellent objects for photomicrography and they provide maximum protection for minute dissected parts, which is an important consideration in the case of primary types particularly. On the other hand, it is sometimes crucially important to retain the possibility of examining the structures from different angles and / or to avoid any kind of distortion from a cover glass. Also, the time expenditure involved in consistent preparation of high-quality slide preparations (see Robinson 1976) can be difficult to justify in those cases where examination of many specimens is desirable or necessary. Here the storage of genitalia preparations in glycerol and associated with the pinned specimens, is often a satisfactory solution. (The availability in recent decades of light-weight plastic vials with silicone rubber stoppers has largely eliminated those misgivings about the vial-storage method, which were justified, when only glass vials with cork stoppers were available.) If certain technical advances become routine requirements in taxonomy, further impositions are likely to be placed on taxonomists. Detailed microscopical observation of many skeletal features (see, e. g., Lee & Brown 2006), SEM examination of scales and antennal sensilla and, increasingly, molecular barcoding are notable examples. Even if taxonomists themselves are not involved in laboratory procedures, the removal, storage and recording of tissue samples are likely to prove to be time consuming. In contrast, other innovations (such as electronic description templates, high-quality imaging, of entire specimens and morphological details including slide mounts) may make conventional procedures less labour-intensive. The net consequences for throughput in revisionary Lepidoptera taxonomy remain to be seen. Fig. 5 (lower graph) reveals a disconcerting observation for the abovementioned tightening of working practices, namely the legacy of synonyms created during phases of unconcerned mass-descriptions. Species concepts. Any discussion of species diversity must in principle make reference to the species concept adopted. If asked, most taxonomists would probably say that their species are inferred to be reproductively isolated, and that morphological distinction is a surrogate measure of reproductive disjunction. In practice, most species are described on the basis of the kind of morphological differences (sometimes informed by natural history observations) traditionally used – many species being described before the biological species concept was articulated. This gives some reassuring constancy to those interested in using taxonomic data for understanding species diversity. Should, however, alternative species concepts be introduced, such as inflating subspecies to the level of species, our assessment of species numbers might change significantly.	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
039C87C3FFD6DC3FFF3EE0EFFD60F9C4.taxon	description	The fossil record of Lepidoptera (as of other insects) was recently reviewed by Grimaldi & Engel (2005), whose excellent account is recommended strongly. The oldest fossil currently believed to belong to the Lepidoptera is Archaeolepis manae from the Lower Jurassic (ca 190 MYO) and according to Grimaldi & 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., Micropterigidae) 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 Heterobathmiidae + Glossata. Following Labandeira et al. (1994), 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 & Engel rightly caution about the uncertainty inherent in identifying leaf miners. In any case there is little doubt that the main radiation of the Lepidoptera 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 Lepidoptera, 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 Pieridae and Nymphalidae as estimated from molecular evolution is more than 70 MYO, hence the butterflies as a whole must be somewhat older (Braby et al. 2006, Wahlberg 2006).	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
039C87C3FFD2DC04FF3EE448FED9FBFA.taxon	description	Regional / local checklists come in many forms. The only major zoogeographical region for which complete, modern Lepidoptera checklists are available are North America (Hodges et al., 1983) and Australia (Nielsen et al. 1996); a checklist of Neotropical Lepidoptera is in the course of publication since 1984 (Heppner ed.). Important sub-regional checklists include, e. g., those by Karsholt & Razowski (1996) on Europe (and a more recent electronic list is available on http: // www. faunaeur. org /), Heppner (1992) on Taiwan and Vári et al. (2002) on Southern Africa. As for other insect orders a useful survey of identification works, including primary research articles, was given by Hollis (1980). Works of regional scope that proved invaluable tools for Lepidoptera systematists worldwide include the accounts of Nearctic Lepidoptera immatures in ' Immature Insects' (F. Stehr ed, 1987), the book by Common (1990) on Australian moths, and Holloway et al. 2001 on Malesian Lepidoptera. The somewhat detailed account of Lepidoptera systematics on a global basis presented in the multi-author treatment of the order in the ' Handbuch der Zoologie / Handbook of Zoology ' series (Kristensen ed. 1998) also gives many references to identification literature. Many important internet resources for systematic Lepidopterology can be found at http: // www. lepsoc. org / lepidoptera _ websites _ both. php. Of particular note are also the principal websites that give access to important classical descriptive literature, thereby greatly facilitating revisionary studies to workers who have no easy access to comprehensive libraries: ‘ Biodiversity Heritage Library’ (http: // www. biodiversitylibrary. org /) and ‘ Animal Base’ (http: // www. animalbase. uni-goettingen. de / zooweb / servlet / AnimalBase / search # about); both are still in development	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
039C87C3FFEDDC07FF3EE5E7FC7AFC62.taxon	description	It is difficult to estimate the number of entomologists presently engaged in research on Lepidoptera systematics. Most professional workers who make contributions in this field have demanding commitments of other kinds as well (teaching, curatorial, managerial) as have, of course, the amateur taxonomists. The figure for full-time research equivalents of the contemporary Lepidoptera systematics workforce is, therefore, very much lower than the number of authors of publications on the subject. The specialist coverage of lepidopteran subgroups is somewhat uneven, and the exceedingly low number of taxonomists working on the most speciesrich families, such as Pyralidae, Crambidae, Geometridae and Noctuidae (Holarctic faunas excepted), is a particular problem. Lepidopterists make no exception to the gloomy picture of the age-structure in the taxonomists' community as a whole, as underscored in the afore-mentioned Gaston & May survey: to our knowledge there are at present disturbingly few Ph. D. students in Lepidoptera systematics worldwide. With so much ground still to be covered, and so few workers to cover it, contemporary systematists working on the largest insect orders should pay considerable attention to the criteria on which research priorities should be set. To those systematists who are employed by primarily non-taxonomically orientated institutions (and they constitute a considerable proportion of the grand total), priorities naturally come in the form of taxa which contain economically important species, or which prove to be convenient target taxa for sampling and monitoring in biodiversity assessment / nature management contexts. Moreover, of course, workers in such institutions are usually supposed to deal with geographically restricted faunas. But what are the guidelines for the few systematists in those universities or major research museums, where licentia academica is still supposed to prevail? Arguably phylogeny-based generic / genus group classifications of higher taxa (family / subfamily level) should be among the highest contemporary priorities. Good examples of such classifications of a global scope of genus-rich taxa are Kitching's (1987) treatment of Noctuidae-Plusiinae and Miller's (1991) of Notodontidae. A number of studies of this kind have been restricted to regional faunas; examples are the treatments of Holarctic sesiids (Naumann 1971 / 1977) and Australian tineids (Robinson & Nielsen 1993), oecophorids (Common 1994, 1997, 2000) and olethreutine tortricids (Horak 2006). In spite of their limitations such regionally based phylogenies may prove useful platforms for developing global classifications. Similarly, purely descriptive, but comprehensive and well-illustrated generic accounts, whether global (e. g., that of Pitkin et al. 2007 on Geometrinae-Pseudoterpini) or regional (e. g., that of Pitkin 2002 on Neotropical Geometridae-Ennominae) are invaluable stepping stones on the way towards the desirable phylogeny-based classifications. Classificatory ' skeletons' of the said kinds are essential for providing appropriate frameworks for specieslevel revisionary taxonomy. Opinions are divided about priority setting in the latter approach, but few biologists will contest that describing all species and classifying them in accordance with their position on " the one true tree of life " is the ultimate goal of systematics. Genuine progress will depend on a balanced partition of resource investments in bottom up projects in the form of species-level revisions, and top down projects in the form of generic / genus group classifications of higher taxa (family / subfamily level). In this respect contemporary systematic lepidopterology is no different from contemporary systematization of other groups of organisms. What is special about systematization of Lepidoptera as well as the other ' megadiverse' insect orders is the sheer magnitude of the task. Development of the afore-mentioned classificatory ' skeletons' of the largest family-group taxa will be undertakings which will require time investment (material gathering, preparations and descriptive / pictorial documentation of morphological characters, gene sequencing, phylogenetic analyses, etc) beyond the capacity of individual researchers, and they will be practicable only to research teams with access to major institutions with truly comprehensive collections. If, therefore, we are to have the much needed globa l classificatory ' skeletons' for e. g., the ' catocaline' or ' hadenine' noctuids, ennomine or larentiine geometrids, phycitine or pyraustine pyraloids etc., etc., then we shall need initiatives to this effect being supported at the managerial level in major research museums, or perhaps more realistically in consortia thereof. Such initiatives would expectedly come in the form of funded projects with X principal investigators, Y postdocs, Z research students, A technicians and B operation money. Projects of this kind have so far not been commonplace in systematic entomology, but they are what progress in several other scientific disciplines is currently dependent on. There is, currently, considerable interest in the role of the internet in taxonomy, although the diversity of approaches suggests an area in a state of rapid evolution and exploration. The semi-structured and formalised nature of taxonomic content lends itself to being transposed into database format, rendering it potentially far more accessible and flexible. But the landscape of taxonomic cyberspace is complex. Taxonomic data on the web fall into a few main categories. Ever more descriptive taxonomy is becoming available online as pdf files through electronic journals or journals, like this one, that are largely electronic. Although digital, the medium here remains, like paper, a fixed one. Fixed medium has the benefit of being archivable and helps protect nomenclatural stability. Contributions can also be quality controlled by the traditional review process. But like paper publications, those in pdf format lack the capacity to be updated other than by further and separate publication of results. By contrast, web-pages posted as HTML are capable of being updated, but the ease by which they can be created and posted on the internet means that quality is both highly variable and typically not indicated to readers. Many such websites for Lepidoptera are accessible on the web. An experiment to provide comprehensive web-based taxonomic revisions in a more controlled environment is the aim of the CATE project (www. cate-project. org), which is being developed currently. One of the two taxa used to trial and demonstrate the CATE method and principles is the Sphingidae (ca 1400 species). Revisions created through the CATE system (see Godfray et al., in press; Scoble et al., in press) are updatable: anyone with access to the web can submit a proposal to a CATE website. Proposals may range from simple geographical records to descriptions of new taxa. However, peer-review and editorial procedures are built into the workflow so that quality control can be maintained. Less structured, but highly updatable, mechanisms for taxonomy are ‘ scratchpads’ (V. Smith, pers. comm.; www. editwebrevisions. info / scratchpadSiteList). These are based on a content management system and are intended to encourage web-based collaboration between taxonomists. The most recent large scale and highly ambitious international project called the Encyclopedia of Life (see www. eol. org) is aimed at building a web-page for each of the ca 1.8 million species. Aggregation (‘ mashup’) technology will be used to source taxonomic information from across the web with the intention of encouraging taxonomists to structure and edit the resulting content. The future of taxonomy looks increasingly as if the internet will eventually predominate as the platform for providing access to information about biodiversity, including all kinds of relevant taxonomic and other data. Software is ever more able to provide users with the data they require in a customised form, such as global or regional checklists, maps of species distributions based on point data, and descriptions of species. The main challenges in this endeavour are generating collaboration among a scattered and variable community, developing the capacity to atomise data to structure underlying databases, and quality control. There is an increasing expectation that information of all kinds should be available on the web. The taxonomic impediment shows little sign of being resolved by more conventional means and the best chance of integrating the effort of all Lepidoptera taxonomists, be they professional or not, is likely to be through collaboration in cyberspace. It will take a new generation of taxonomists and technology (that of the semantic web particularly) for this vision to be realised, but it is more likely to bear fruit than a more dirigist approach, which has largely failed to produce an organised and structured way of doing taxonomy. Information and Communication Technologies (ICTs) do not merely provide a more accessible and flexible medium for posting data. Rather they provide a highly accessible (virtual) environment for community involvement by anyone with an internet connection. ICTs profoundly change the collaborative and managerial landscape for improving taxonomic coverage. Optimizing the outcome of this innovation will require a judicious balance between initiatives coming through bottom up engagement by practising taxonomists (of whatever status) and from top down priority setting by management within the systematists' community. Two final points concerning taxonomic research strategy should be mentioned as particularly pertinent to lepidopterology because of its sizeable amateur contingent. The role non-professionals play and continue to play in Lepidoptera taxonomy cannot be overstated. Duckworth, Genoways and Rose (1993) estimated that there were around 2.5 billion specimens (of all taxa) in natural history collections across the world. (L. Speers in Scoble 2003 suggested a range of 1.5 to 3 billion.) Being such a popular group, the Lepidoptera component of this huge estimate is surely significant. Amateurs have contributed to this collective resource in two ways. First, they have supplied institutions with specimens to such an extent that many great holdings would be bereft without the material contributions they have made over more than two and a half centuries. And amateurs follow the tradition and continue to gift material. Second, many specimens continue to be held in private collections to this day — some such holdings competing in terms of importance with collections in state-owned collections. Furthermore, most amateur lepidopterists are not just collectors, but also knowledge creators through their records and publications — publications that are often written in collaboration with professional workers. Amateurs form, then, a significant component of our collective taxonomic knowledge base. Given the popularity of Lepidoptera, their contributions are conspicuously high when we look across all taxa. Two concerns deserve attention. One is that institutions need to be prepared to cope with substantial amounts of material when they are gifted or bequeathed. Storage space and curatorial effort to deal with large amounts of material need careful planning. Such planning includes the capacity to finance the requirements. A second problem arises when provision has not been made for a private collection when the owner dies. Beneficiaries may not share the same concern for such a collection or may fail to understand its scientific value. It is by no means unknown for collections to be broken up and sold on the open market to maximise their financial returns. The issue of stewardship is a responsibility for the lepidopterist community at large — amateur, professional and institutional. As noted above, the rate of description of new taxa by professional taxonomists broadly has decreased as the result of adoption of stricter working standards and the emphasis on revisionary taxonomy. However, new species continue to be described without peer-review and without being placed in a revisionary context. It is a major challenge to professional lepidopterists in leading institutions, which are comparatively well-resourced, to establish close links with the communities of amateur lepidopterists, and less privileged professionals, to optimize the utilization of the considerable labour potential in these communities. If we are to understand and describe, to effect, the diversity of the Lepidoptera, all lepidopterists need to unite in a common effort. Given the number of enthusiasts, these goals are likely to be within our reach.	en	Kristensen, Niels P., Scoble, Malcolm J., Karsholt, Ole (2007): Lepidoptera phylogeny and systematics: the state of inventorying moth and butterfly diversity. Zootaxa 1668: 699-747, DOI: 10.5281/zenodo.274044
