ISSN 0013-8738, Entomological Review, 2012, Vol. 92, No. 4, pp. 390–402. © Pleiades Publishing, Inc., 2012. Original Russian Text © V.G. Mironov, 2012, published in Entomologicheskoe Obozrenie, 2012, Vol. 91, No. 1, pp. 143–159.
Pugs of the Tribe Eupitheciini (Lepidoptera, Geometridae): the History of Study and Distribution over Biogeographic Regions V. G. Mironov Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia Received July 4, 2011
Abstract—The World fauna of the tribe Eupitheciini is the most species-rich in the family Geometridae. This tribe includes about 1900 species (almost 3000 species-group names) from 47 genera; about one third of the genera (15) are monotypic. The generic diversity of Eupitheciini is the highest in the Australian (38 genera, 11 of them endemic) and Oriental regions (32 genera, 4 endemic) and the lowest in the Neotropical Region (possibly one genus only). The faunas of different biogeographic regions can be arranged in following order by their species richness: the Palaearctic (487 species), Oriental (397), Neotropical (346), Australian (251), Afrotropical (198), and Nearctic Regions (166 species). Eupithecia is the most species-rich genus in the family Geometridae and the entire order Lepidoptera, and one of the largest genera in the whole World fauna of insects. The greatest number of species of this genus is recorded in the Palaearctic Region (466 species), where Eupithecia accounts for about 95% of the tribe Eupitheciini. The mainland of the Oriental Region (especially the Himalayas) is also very species-rich; however the proportion of the Eupithecia representatives decreases towards Malaysia, Sundaland, and the Australian Region (about 2% of the tribe). The Eupitheciini faunas have the greatest similarity at the generic level between the Oriental and Australian Regions (the Jaccard and Sørensen coefficient values being 0.62 and 0.77, respectively). The Palaearctic fauna is more similar to the Afrotropical and Oriental faunas at the genus-group level. On the whole, the fauna of the Nearctic Region is similar to that the West Palaearctic, with the exception of the fact that representatives of the genera Gymnoscelis and Chloroclystis are absent in North America, although two endemic genera Nasusina and Prorella are present. At the genus-group level, the Nearctic fauna of Eupitheciini is more similar to the Neotropical (the Jaccard and Sørensen coefficients 0.20 and 0.33, respectively) than to the Palaearctic fauna (0.17 and 0.29). The number of synonymies is very high in the tribe Eupitheciini because of the homogeneity of this group, whose species are difficult to identify without the use of elaborate anatomical techniques. Modern revisions, catalogues, surveys, and atlases on Eupitheciini are absent for many countries and large geographic regions. Revisions of pugs of the tribe Eupitheciini for some biogeographic regions are extremely difficult because of fragmentation of entomological collections including the type specimens of many species-group taxa. A large fraction of synonyms is characteristic of parts of the World with the best known faunas: Europe (64% of synonyms) and North America (39%). On the contrary, the lowest levels of synonymy are typical of the less known faunas of the regions situated at the equatorial latitudes, namely the Neotropical (9%) and Afrotropical (8%) ones. DOI: 10.1134/S0013873812040045
Geometridae is one of the largest lepidopteran families, second in species diversity only to Noctuidae and Pyralidae. This family is widely distributed in all the biogeographic regions and counts from 21000 to 23000 species in the world fauna, according to the recent data of different authors (Heppner, 1991; Gaston et al., 1995; Scoble et al., 1995, 1999). At present, dozens of specialists in many countries are occupied with the study of this group. Numerous revisions of genera, tribes, and even subfamilies in some regions, a great number of descriptions of new species, data from molecular or functional-morphological studies of
many taxa, and synonymization of some old names have resulted in considerably improved knowledge of the taxonomy and distribution of Geometridae. This is presently one of the well studied lepidopteran families, especially at the level of large taxa of the subfamily and tribe rank. A special place among all the geometrid moths belongs to the so-called “pugs” (Blütenspanner) of the tribe Eupitheciini, one of the most diverse and taxonomically difficult groups. The tribe unites representatives of 47 genera almost a third (15) of which are
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monotypic. In its species diversity this tribe has no equals in the family. Almost 1900 species of pugs were described from all the biogeographic regions, which is about one-tenth of the species composition of the family. The type genus of the tribe, Eupithecia Curtis, 1825, is not only the richest in species in the order Lepidoptera but also one of the largest among all the insects. During the last 250 years, over 2200 species-group taxa have been described in it, and almost 1400 species names are still valid. Publications using statistical processing of taxonomic data to estimate the species richness of geometrid moths of different subfamilies and in different regions of the world have appeared only recently (Gaston et al., 1995; Scoble et al., 1995). In the works of this kind, pugs of the tribe Eupitheciini were considered only within the subfamily Larentiinae. However, both by the total number of the taxa described and by the number of valid species names in the world fauna, this tribe is quite comparable with the subfamilies Oenochrominae, Geometrinae, and Sterrhinae (Geometridae) as well as with many other families and large subfamilies of Lepidoptera (Heppner, 1991). In the present paper we have tried to evaluate the degree of knowledge of pugs of the tribe Eupitheciini, compare their diversity at the genus and species levels, and characterize more precisely their distribution and endemism in different biogeographic regions. MATERIALS AND METHODS The study is based on the global database of the world fauna of geometrid moths maintained at the Natural History Museum in London and published as a two-volume catalogue of the family Geometridae (Scoble et al., 1999). This catalogue includes a great body of information: valid names of genera, species and subspecies, synonyms, authorship and year of description, a complete primary bibliography, data on each type series, place of its storage, and the type locality of each particular taxon. During the last 25 years we have studied material (including types) on pugs of the tribe Eupitheciini in the Natural History Museum (London), the Zoological Museums of Munich and Karlsruhe (Germany), Alexander Kӧnig Research Museum (Bonn), the Hungarian Natural History Museum (Budapest), the Zoological Institute of the Russian Academy of Sciences (St. Petersburg), Humboldt Museum of Natural History (Berlin), the Royal Museum of Natural History ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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(Stockholm), the Natural History Museum (Vienna), the Agricultural Institute of Estonia (Tartu), the Institute for Zoology of the Chinese Academy of Sciences (Beijing), the zoological museums of Copenhagen, Moscow, and Kiev Universities, and also private collections of P. Skou (Stenstrup, Denmark), M. Fibiger (Sorø, Denmark), Dr. U. Ratzel (Karlsruhe, Germany), M. Sommerer (Munich, Germany), Dr. D. Laslo and O. Pekarsky (Budapest, Hungary). As the result of these studies it was possible to reveal some gaps in the Catalogue (Scoble et al., 1999) and obtain new, more complete data from two adjoining biogeographic regions: the Palaearctic and Oriental ones. Data from the Palaearctic are reflected in publications embracing the faunas of Europe (Mironov, 2003), the Canary Islands (Bacallado and Mironov, 2003), the Russian Far East (Mironov, 2005), the Western Himalayas (Mironov et al., 2008a– 2008c), China (Mironov et al., 2004a–2004d, 2006, 2011), Japan (Mironov and Galsworthy, 2010b), and Mongolia (Mironov et al., in litt.). For the Oriental Region, material from Southeast Asia (Galsworthy and Mironov, 2005; Mironov and Galsworthy, 2009a, 2009b), Taiwan (Mironov and Galsworthy, 2007), Nepal, India, and Sri Lanka (Mironov and Galsworthy, 2010a) were studied. In the analysis of taxonomic data, recent publications on all the biogeographic regions, particularly the Oriental and Australian ones, were taken into account (Galsworthy, 1999, 2003; Inoue, 2000, 2002). Altogether, during the 12 years which elapsed after the publication of the Catalogue (Scoble et al., 1999), 196 new species and 6 subspecies of Eupitheciini have been described; names of 8 species-group taxa have been restored from synonyms, 4 taxa elevated to species rank; 183 species-group names from different genera of the tribe have been synonymized for the first time, and 2 names have been reduced to subspecies. The most important taxonomic changes involved the faunas of the Palaearctic and Oriental Regions. The knowledge of the species composition of these faunas and ranges of individual species was considerably extended. For each biogeographic region, only the taxa described from that region were considered. The total number of species-group taxa includes all the known species and subspecies as well as varieties (variants) and forms described before 1961 and possessing the species group rank according to Articles 15.2 and
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45.6.4 of the International Code of Zoological Nomenclature (2000). The genus- and species-group replacement names and the scarce new combinations within the tribe Eupitheciini were not taken into account since they did not affect the total number of valid names of genera and species or the number of synonyms. The only exceptions were taxa which received a new, higher rank (subspecies elevated to species), taxa whose names were restored from synonyms, and also taxa transferred to other tribes of the family Geometridae. The species names whose rank had been reduced to subspecies were treated as synonymized, since the number of valid species names was thus reduced. The faunistic similarity of different biogeographic regions at the genus level was assessed using the common Jaccard and Sørensen coefficients. RESULTS AND DISCUSSION The first species of pugs (Phalaena Geometra rectangulata and Phalaena Geometra succenturiata) were described by Carl Linnaeus in his Systema Naturae (Linnaeus, 1758). There was little taxonomic activity throughout the rest of the XVIII century after publication of the book till the first decade of the XIX century. During this period only 26 species were described from Europe, of which 14 names were later reduced to synonyms. The reason for this was lack of coordination between European naturalists who worked at that time in Sweden, Denmark, Germany, Austria, France, and Italy, lack and inaccessibility of some publications, incompleteness of original descriptions, paucity and sometimes low quality of illustrations, and sometimes their complete absence. Description of new species of geometrid moths from Europe including those placed in future into the tribe Eupitheciini began in the first half of the XIX century with publications of Jacob Hübner (1796– 1838) in Germany and Adrian Haworth (1803–1928) in Britain. These authors described 38 taxa of pugs half of which were subsequently synonymized. The first species from the territory of the Russian Empire, Acidalia extensaria, was described only in 1845 (Freyer, 1845). By the early XX century, owing to the efforts of many scientists and naturalists, the species composition of the European fauna of pugs was roughly outlined; 102 species were described, comprising 77% of the presently known number; 150 more species names were synonymized during that time and later.
Active description of new species from other continents started in the second half of the XIX century. The greatest number of new descriptions of speciesgroup taxa appeared in the last decade of the XIX century and the first decade of the XX century (Fig. 1); they were mostly made by specialists of two main European schools, the English and the German one. The first known species of pugs were given trinomial names since Linnaeus included them into the genus Phalaena Geometra. Subsequent authors already used the strict binomial nomenclature. For almost a century, beginning with 1804, new taxa of pugs were described in several genera: Larentia Treitschke, 1825; Cidaria Treitschke, 1825; Acidalia Treitschke, 1825; Melanthia Duponchel, 1829. Due to their small size, some species were placed in the genera Tinea Linnaeus, 1758 (Tineidae) and Pyralis Linnaeus, 1758 (Pyralidae). The name Eupithecia was introduced for the first time by John Curtis (1825) who included 38 species of British pugs into this genus. Somewhat later, according to the chronological data of Hemming (1937), Hübner (1816–1826) described a great number of new genera of Geometridae and included 31 species of pugs into 7 of them: Tephroclystia Hübner, 1825 (10 species); Dyscymatoge Hübner, 1825 (7); Tarachia Hübner, 1825 (3); Leucocora Hübner, 1825 (4); Arcyonia Hübner, 1825 (2); Eucymatoge Hübner, 1825 (3); and Hypepirritis Hübner, 1825 (2 species). All these genera were established on the sole basis of wing coloration and pattern. During the period between 1839 and 1999, 21 genera were described with type species from among the representatives of the genus Eupithecia. Only 4 of them are still valid: Ardonis Moore, 1888, Dissolophodes Warren, 1907, Casuariclystis Holloway, 1997, and Sigilliclystis Galsworthy, 1999. As with species, the greatest number of genera was described at the turn of the XX century, as the result of processing vast materials from different biogeographic regions kept at the British Museum. Priority in this certainly belongs to William Warren (1895, 1903, 1906, 1907, etc.) who described almost half (46%) of the genera of the tribe Eupitheciini. The faunas of pugs in each biogeographic region have specific traits, differing in the richness of genera, species and species groups, in the level of endemism, and also in the history of study, the degree of exploration, the fraction of synonymized names, etc. The Palaearctic Region (31 027 mln km2). This is one of the best studied regions of the globe. Due to its ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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Fig. 1. The dynamics of description of the species-group taxa of the tribe Eupitheciini in different periods. Abscissa, years; ordinate, the number of the species described. Black circles, the number of valid names; light circles, the general number of names.
vast territory extending over great distances in longitude and latitude, the presence of a great number of natural zones, and highly partitioned relief with several isolated high mountain systems, this region certainly has the most diverse species composition of Eupitheciini. The study of the Palaearctic fauna has a rich history of more than 250 years. Among a vast number of publications on pugs, the most important monographs of Karl Dietze (1910, 1913) and Louis Prout (1912–1916, 1934–1954) should be mentioned which hold good even at present. These monographs include descriptions, distributional data, and a great number of photographs (including those of type specimens) or drawings of the great majority of species, subspecies, forms, and variants of Eupitheciini known at that time from the Palaearctic. Although the Palaearctic pugs were described by nearly 150 scientists and naturalists, 90 (or 60%) of them published only one or two species names. Before 2011, 1057 taxa of the species rank had been described, including 180 subspecies, forms, and variants; 388 names (almost 45%) were subsequently synonymized. This level of synonymy is the highest among Eupitheciini of all the biogeographic regions (Fig. 3). On the whole, the Palaearctic fauna counts 487 species of pugs from 19 genera. The overwhelming majority of them (460 species, or 95%) belong to the genus Eupithecia. The most fully studied is the fauna of Europe counting 4 genera and 135 species of which 114 species (almost 85%) were described from this territory; 240 (64%) of species names were synonymized by different authors. The genera Gymnoscelis Mabille, 1868, Chloroclystis Hübner, 1825, and Pasiphila Meyrick, 1883 are represented in the EuroENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
pean fauna by a few species whose ranges embrace vast territories, extending as far eastwards as Mongolia and even Japan and the south provinces of China. All of them may have originated from the Oriental Region where the above genera are represented by numerous species. Within the genus Eupithecia, the most diverse in Europe are the species groups interruptofasciata (14 species), egenaria, satyrata, and semigraphata (9 species each) which mainly include species of the Mediterranean origin. A total of 44 species from the genera Gymnoscelis and Eupithecia were described from North Africa and adjoining Macaronesian islands (the Azores, Madeira, Canary and Cape Verde islands); 38% of the names were synonymized. Most species belong to the genus Eupithecia which is mostly represented there by the Mediterranean and Atlantic xerophilous species of the groups interruptofasciata (11 species), egenaria (9), and centaureata (6 species). The Asian part of the Palaearctic contains the richest and most diverse fauna of Eupitheciini. Almost 330 species of pugs from 7 genera were described from this territory, and 177 more names of the species rank (35%) were synonymized. In the fauna of West and Central Asia the greatest diversity is to be found in the mainly xerophilous species groups of the genus Eupithecia: venosata (20 species), graphata (19), innotata and gueneata (14 each), centaureata (12), and ultimaria (9). Pugs of these groups develop on umbellate, carnation, composite, and tamarix plants well represented in the flora of West and Central Asia. Similar to Europe, only few representatives of the genera Gymnoscelis, Chloroclystis, and Pasiphila occur there.
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Fig. 2. Growth of the number of the species-group taxa described from the tribe Eupitheciini in different geographical regions. Abscissa, years; ordinate, the number of the species described. Black circles, the number of valid names; light circles, the general number of names. AUS, Australian; AFR, Afrotropical; NEA, Nearctic; NEO, Neotropical; ORI, Oriental; PAL, Palaearctic Region.
The East Asian part of the Palaearctic Region is characterized by a high diversity of genera. Besides 4 genera mentioned for the west part of the Palaearctic, the East Asian fauna includes representatives of Axinoptera Hampson, 1893, Mesoptila Meyrick, 1891, and Spiralisigna Holloway, 1997, which are of Oriental origin. In the genus Eupithecia, the groups haworthiata (11 species), undata (10), egenaria (14), lanceata (8), satyrata (14), and subfuscata (8 species) are especially diverse. Some representatives of typi-
cally Oriental species groups of this genus, such as proterva, propagata, and rajata also occur there, reaching the southeast regions of the Palaearctic, and some of them extend even to South Primorye and the Hokkaido Island. During the last decade considerable progress has been attained in the study of the pug fauna of the Palaearctic region. Many names have been reduced to synonyms but the number of new descriptions is even greater, which can be clearly seen in Fig. 2. ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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The Nearctic Region (14 724 mln km2). The study of pugs of the tribe Eupitheciini in North America started a century later than in Europe. The two first species from the Nearctic were described by Achille Guinée (1858), followed by a series of new descriptions by H.B. Mӧschler, F. Walker, K. Dietze, and A.S. Packard. Most taxa of the species rank (207 names out of 268, or 78%) were described during 50 years from 1896 till the end of the World War II (Fig. 2), mainly by G.D. Hulst, R.F. Pearsall, S.E. Cassino, L.W. Swett, and J.H. McDunnough. The publications of the above authors containing the original descriptions are not cited here since this would take up too much space. The pug fauna of the Nearctic is relatively poor and unvaried. Only 4 genera of the tribe are represented in North America of which 2 are endemic and quite small (Nasusina Pearsall, 1908 and Prorella Barnes et McDunnough, 1918), whereas the genus Pasiphila is represented by a single species probably introduced from Europe (Ferguson, 1972). The Nearctic fauna is generally similar to the Palaearctic one. Besides the 11 species shared by these biogeographic regions, the faunas of the Nearctic and the Palaearctic include 23 common species groups of the genus Eupithecia (Mironov, 2003). Representatives of only 3 species groups, namely palpata, stellata, and annulata, are endemic to the New World. Five pairs of sibling species from the Nearctic and the Palaearctic, very similar both in habitus and the genital structure, provide very interesting objects of molecular studies: Eu. johnstoni McDunnough, 1945 and Eu. linariata ([Denis et Schiffermüller], 1775); Eu. misturata (Hulst, 1896) and Eu. silenata Assmann, 1848; Eu. albicapitata Packard, 1876 and Eu. analoga Djakonov, 1926; Eu. mutata Pearsall, 1908 and Eu. abietaria (Goeze, 1781); Eu. niphadophilata (Dyar, 1904) and Eu. pusillata ([Denis et Schiffermüller], 1775). The greatest species diversity and specificity of the Nearctic fauna is characteristic of the western mountainous part of North America: the Rocky Mountains, the south of the USA and north of Mexico. Of 175 species of pugs known from the Nearctic Region, 166 (95%) were described from North America, 164 being endemic to the continent. The fraction of synonymized names of Eupitheciini is 39%, which is one of the largest as compared with the other biogeographic regions (Fig. 3). There are several reasons for it. On the one hand, poor knowledge of the European fauna by American entomologists (22 synonymized names) and numerous parallel descriptions ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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Fig. 3. The ratio between the number of valid names and the number of synonyms in different biogeographical regions.
made between 1904 and 1912 (53 synonyms) played a certain role. On the other hand, the Nearctic fauna of Eupitheciini was repeatedly revised at a sufficiently high level (McDunnough, 1949; Ferguson, 1983; Blanchard et Knudson, 1985; Bolte, 1990) and is presently well studied. The Neotropical Region (12 027 mln km2). The pug fauna of this region is extremely rich; according to the latest data, it is represented by 346 species, almost all of them from the genus Eupithecia. The first species from the Neotropical Region were described by Francis Walker (1863) and Pieter Snellen (1874) based on material from Venezuela, Honduras, and Colombia. However, the great majority of taxa from Central and South America were described later, at the turn of the XX century, mainly by W. Warren who processed the huge collections in the Natural History Museum in London. A real boom in new descriptions of Eupitheciini from the Neotropical Region fell on the two last decades of the XX century (Fig. 2). On the whole, 70% of all the pug species known from this region were described by five authors: P. Dognin, W. Warren, W. Schaus, C. Herbulot, and F. Rindge. A very small fraction of synonyms (9%) in the Neotropical fauna can be explained by the absence of the regional revision of the genus Eupithecia (Fig. 3). A revision of the Neotropical Eupitheciini would be a difficult task since the type material is scattered over different museums of Europe (London, Munich, Frankfurt-on-Main, Munchenberg, Copenhagen, Brussels) and America (Washington, New York, Ottawa), as well as those of Pretoria (South Africa), and San Miguel de Tucumán (Argentina). Claude Herbulot (2001) gave the exact number of the known species of this genus broken down by the countries from which they had been described, and united them into 7 groups. The greatest number of
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species (149) falls on the group distributed in 4 countries (Colombia, Ecuador, Peru, and Bolivia), through whose territories the Andes stretch. However, the mostly mountain fauna of Chile was distinguished as a separate group without special explanations. According to the data of Herbulot (1977), 15 species of Eupithecia occur on the east slopes of the Andes about 3900 m above sea level, which makes up 50% of all the pugs, or 30% of all the known lepidopterans inhabiting these alpine areas. With the altitude decreasing to 2400 m, the number of species increases but their fraction in the number of all the Geometridae and Lepidoptera decreases to 15 and 5%, respectively. Brehm and Fieder (2003) explain the species richness of Eupithecia in the Andean highlands by the diversity of composites which are the probable food plants of the larvae. The males of many species of Eupithecia in the Neotropical Region reveal a surprising diversity in the antennal morphology, the hind wing shape, and especially in the position of androconial structures on the fore and hind wings (Herbulot, 1987). However, the genital morphology of both sexes indicates the presence of few species groups in this genus. In particular, only 4 sections of allied species with a small range of variation within each of them were distinguished among 47 taxa of Eupithecia from Chile and the Juan Fernández Islands (Rindge, 1987). The relatively uniform genital structure of many species may be the result of the recent origin of the fauna of Eupitheciini of Central and South America. The most pronounced endemism is characteristic of the Galapagos Islands which are remote from the continent. Among 13 species of Geometridae known from these islands, 3 very small species of the genus Eupithecia are endemic to the archipelago (Rindge, 1973; Landry et Rindge, 1995). The similarity of their habitus is deceptive; considerable differences in the genital structure, especially in females indicate that these species belong to two morphologically distant groups. The Afrotropical (Ethiopian) Region (16 359 mln km2). The Afrotropical Region is dominated by representatives of the subfamily Ennominae, with a total of 1608 species recorded (Herbulot, 1992). The subfamily Larentiinae is three times less diverse in the region: it includes 467 species, almost half of which are pugs. The first species of Eupitheciini were described by Guenée (1858) and Walker (1862) from the former
British colonies, South Africa and Sierra-Leone. Most of the pug taxa from the Afrotropical Region were, however, described throughout several decades in the middle of the XX century (Fig. 2). Among 18 authors of species names, almost 80% of the species were described by 4 authors: W. Warren, L.B. Prout, D.S. Fletcher, and C. Herbulot. About a quarter of the species is known from the former French colonies: Madagascar and the neighboring small islands in the Indian Ocean (Comoros, Seychelles, and Mascarenes). Some authors consider the Madagascar fauna separately from the continental Afrotropical one, noting its extreme endemism. According to our data, 20 of 31 species of Eupitheciini, or 65% occur only on Madagascar, whereas the fraction of endemics rises to 77% if the species shared with the Comoros are taken into account. Nevertheless, in spite of the high endemism level, I am inclined to agree with Scoble and coauthors (1995) that the Madagascar fauna should not be separated from the Afrotropical one. First, almost a quarter of the Eupitheciini species present in Madagascar also occur in East and South Africa; second, most of the species endemic to Madagascar have close relatives on the African continent. In addition, 2 species: Eupithecia sporobola Herbulot, 1988 and E. tranquilla Herbulot, 1988, are closely related to members of the group haworthiata described from the Comoros, Kenya, and Zimbabwe, and also to species of this group widespread in the Oriental Region. Altogether, 198 species of Eupitheciini are described from the Afrotropical Region, 143 of them (more than 70% of all the pugs) belonging to the genus Eupithecia. At least 10 species groups of this genus are represented in the region, of which 8 are shared with the Palaearctic and some, also with the Nearctic and Oriental regions. The groups haworthiata, egenaria, innotata, and russeliata can be named among the latter. Especially rich is the endemic group dilucida (not less than 13 species). Besides Eupithecia, the Afrotropical fauna includes representatives of the genera Casuariclystis Holloway, 1997 (1 species), Gymnoscelis Mabille, 1868 (13 species), and Mesocolpia Warren, 1901 (8 species). The generic identity of 32 species has not been reliably determined. To date, the most fully studied is the fauna of the south part of the African continent including South Africa, Lesotho, Swaziland, Namibia, Botswana, Zimbabwe, and Mozambique (Janse, 1933–1935; Vári et al., 2002). The tribe Eupitheciini is represented by 61 species in this large territory. ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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In spite of the great number of publications, the pug fauna not only of the Afrotropical Region but also of the whole of Africa needs further investigation, especially the zone of tropical rain forests and the west part of the continent north of the Equator. The need for revision is indirectly supported by the low fraction (8%) of synonymized names of pugs in this region (Fig. 3). The Oriental Region (6659 mln km2). The pug fauna of this comparatively small biogeographic region is one of the richest, second only to that of the Palaearctic. Starting with Guenée (1858), the author of the first species from India (Eupithecia rajata), 397 species of 32 genera of Eupitheciini were described from the Oriental Region. Most of the speciesgroup taxa became known owing to the works of British entomologists (see Fig. 2), almost 54% of the species and subspecies being described by L.B. Prout, W. Warren, F. Walker, and G.F. Hampson. The monotypic genera Dasimatia Warren, 1898, Eva Vojnits, 1981, Eupithystis Holloway, 1997, and also the genus Celaenaclystis Holloway, 1997 with 2 species are endemic to the Oriental Region, yet half of its fauna is formed by 200 species of the genus Eupithecia. Almost all the Palaearctic species groups of this genus are also represented in the Oriental Region. The greatest species diversity is characteristic of the groups haworthiata, tripunctaria, egenaria, and especially proterva, propagata, and rajata, so that the above groups may be of Oriental origin. This is indicated not only by the level of species and morphological diversity but also by the presence of species with plesiomorphic character state in the genital structure. It was in this biogeographic region, in East India (West Bengal and Sikkim) that a species was found combining characters of the groups haworthiata and undata, which differ in the type of uncus (Mironov and Galsworthy, 2010a). The small group kamburonga with 6 species, also distinguished by a peculiar type of uncus, is represented only in the Oriental Region. The species diversity of the genus Eupithecia in Southeast Asia decreases abruptly towards the Malay Peninsula, the Philippines and the Australian Region. In the continental part of the region, especially in the Himalayas, members of this genus comprise more than half of all the pug species. According to the latest data, only 5 species of Eupithecia from different groups occur on the largest and the most fully studied Borneo ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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Island (Holloway, 1997), comprising about 5.4% of the insular fauna of Eupitheciini. The fraction of synonymized species-group names in the Oriental Region is 19% (Fig. 3). This relatively low value can be accounted for by the fact that almost 80% of type species of Eupitheciini are kept in the collections of the British Museum, which also include the great majority of the pug species and subspecies from the Oriental Fauna. The species diversity in the Oriental Region is difficult to assess due to the indistinct boundary with the Palaearctic, namely the presence of the so-called transitional zone (Müller, 1974) between these biogeographic regions. High-altitude regions with a complex relief are characterized by the penetration of the Palaearctic species far to the south, and the Oriental ones, to the north. For example, the fauna of Eupithecia of the West Himalayas consists of Palaearctic species by one third (34–36%), with almost the same fractions of Oriental species and those endemic to this territory. According to our data, the pug faunas of Nepal and the south provinces of China also include a great number of alpine Palaearctic species. Taking into account the climate and flora this is not surprising. According to the botanical studies, the subalpine and alpine vegetation extends up to 4500–5000 m above sea level on the southwest slopes of the High Himalayas, whereas the temperate forests growing lower include typical trees and shrubs of the temperate latitudes: Abies, Picea, Pinus, Juniperus, Betula, Acer, Sorbus, Berberis, Rhododendron, etc. The boundary between the Palaearctic and the Oriental Regions in the High Himalayas is conventionally drawn at about 3000 m above sea level. The mixed fauna of the Palaearctic and Oriental species is also typical of Taiwan. Among the 46 species of Eupithecia known from this tropical island, there are equal fractions (23.5% each) of Palaearctic and Oriental species, the remaining 53% being endemic to Taiwan; their ancestors must have migrated to the island not only from the south but also from the north continental regions of China and from the Japanese Islands. The boundary between the Oriental and the Australian regions is also ambiguous. At least 5 main variants of delimiting these regions are known (Müller, 1974). In my opinion, for pugs this boundary should be drawn not along the better known “Weber’s Line” but rather along the “Lydekker’s Line” separating the
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Sunda and Maluku Islands from New Guinea. Thus, the Eupitheciini fauna of Sulawesi Island includes about 70% species shared with the rest of the Sunda Islands and partly with Southeast Asia, whereas it has less than 50% of species in common with the Australian Region, or to be more precise, with New Guinea and Australia. The Australian Region (5870 mln km2). The first species of this region were described by Guenée (1858) and Walker (1862), mainly from Australia and New Zealand. Most of the species were described during the last decade of the XIX century and the first half of XX century (Fig. 2), until the last publication of Prout (1958). More than 80% of pug species were described by the British and Australian experts W. Warren, L.B. Prout, E. Meyrick, and A.J. Turner. To date, 251 species of 38 genera of Eupitheciini have been described in Australia. The most diverse genera are Tripteridia Warren, 1903 (35 species), Gymnoscelis Mabille, 1868 (30 species), and Pasiphila Meyrick, 1883 (28 species), the latter being mainly recorded in New Zealand. Only 20 species presently remain in the genus Eupithecia. The fraction of synonymized species-group names is relatively low (17%), which can be clearly seen in Fig. 3. There are few species shared with the Oriental Region; Holloway (1997) listed only 26 species of Eupitheciini, the characteristic inhabitants of the tropical forests of the “Indo-Australian” region, which are mainly common for Borneo and New Guinea. The region is characterized by a large fraction of endemic genera and species of Eupitheciini: 11 genera are endemic, of which 9 are monotypic. The greatest number of endemic genera (7) is in New Guinea, and the smallest number is in Australia (3) and New Zealand (1 genus). The Eupitheciini of the Hawaii, the most distant of the Pacific islands, are completely endemic. Meyrick (1899) assumed the Asian origin of Eupithecia occurring in the Hawaii but his opinion does not agree with our data. Since not a single close representative of this tribe exists in the Oriental Region, the ancestors of the Hawaiian pugs should be rather looked for in Australia or the equatorial Pacific islands. There is unique similarity, at least in external characters, between some Australian species, such as “Eupithecia” filata Guenée, 1858, and the Hawaiian Eupithecia staurophragma Meyrick, 1899. The structure of genitalia in the above species also shows them to be distantly related. It can be only supposed that the
numerous Melanesian and Micronesian islands served as stepping stones during dispersal of Eupitheciini throughout the Pacific, from Australia to the Hawaii. The fauna of Australia and adjacent islands was studied for over 50 years by A.J. Turner who described 48 species, i.e., almost 65% of the pugs of this region. In spite of the works of Turner and some other summary publications on the faunas of Australia (McQuillan and Edwards, 1996) and New Zealand (Dugdale, 1988), the pugs of this biogeographic region are still insufficiently studied. The taxonomic position of many species remains obscure while some genera need revision. The studies are hampered by the fact that most type specimens are divided between the British Museum and the Australian National Insect Collection in Canberra. The species diversity of pugs varies considerably in different biogeographic regions of the Earth. This group is the most diverse in the Palaearctic, Oriental, and Neotropical Regions (Table 1). As compared to the total number of species in the subfamily Larentiinae, the fraction of Eupitheciini is noticeably greater in the Palaearctic and Australian Regions but is nearly half as great in the fauna of the Neotropical Region, where the total number of species in the genera Eois Hübner, 1818 and Psaliodes Guenée, 1858 is not less than that of pugs, while representatives of the subfamily Ennominae prevail in species diversity. The borders of ranges of many pug species remain uncertain. Many species are only known by occasional findings, sometimes even by a single specimen from the type locality. The territories on the boundary between biogeographic regions, characterized by mixing of representatives of different biotas, are not sufficiently studied; therefore analysis of the similarity of faunas at the species level has not yet been possible. In this communication, analysis of similarity of the Eupitheciini faunas of different biogeographic regions was performed at the genus level, using Jaccard and Sørensen coefficients. The results of this analysis are given in Table 2. As expected, the faunas of the Oriental and Australian regions are the most similar ones, the Jaccard and Sørensen coefficients being 0.62 and 0.77, respectively. The Palaearctic fauna is closer to the Afrotropical and Oriental faunas in the composition of the pug genera. The poor Nearctic fauna of Eupitheciini, in spite of a number of species being shared with the Palaearctic, differs from the latter in its generic comENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
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Table 1. Comparative species diversity of the family Geometridae in different biogeographic regions Biogeographic regions
Taxa Family Geometridae Subfamily Archiearinae Subfamily Ennominae Subfamily Geometrinae Subfamily Larentiinae Tribe Eupitheciini Subfamily Oenochrominae Subfamily Sterrhinae
PAL
NEA
NEO
AFR
ORI
AUS
0.16 0.40 0.13 0.10 0.21
0.06 0.20 0.07 0.03 0.08
0.32 0.40 0.36 0.20 0.31
0.15 0 0.16 0.25 0.08
0.25 0 0.23 0.36 0.24
0.06 0 0.05 0.06 0.08
0.26 0.05 0.18
0.09 0.01 0.03
0.18 0.14 0.33
0.11 0.07 0.18
0.22 0.30 0.23
0.14 0.43 0.05
Notes: PAL, Palaearctic; NEA, Nearctic; NEO, Neotropical; AFR, Afrotropical; ORI, Oriental; AUS, Australian.
Table 2. Similarity of the generic composition of the faunas of the tribe Eupitheciini of different biogeographic regions: Jaccard and Sørensen (in brackets) coefficients Biogeographic regions
PAL
NEA
NEO
AFR
ORI
AUS
PAL
0.17 (0.29)
0.09 (0.17)
0.31 (0.47)
0.23 (0.37)
0.16 (0.28)
NEA
– – 0.17 (0.29)
– –
0.20 (0.33)
0.10 (0.18)
0.06 (0.11)
0.05 (0.09)
NEO
0.09 (0.17)
0.20 (0.33)
0.13 (0.22)
0.18 (0.11)
0.05 (0.10)
AFR
0.31 (0.47)
0.10 (0.18)
– – 0.13 (0.22)
0.18 (0.30)
0.15 (0.26)
ORI
0.23 (0.37)
0.06 (0.11)
0.18 (0.11)
– – 0.18 (0.30)
– –
0.62 (0.77)
AUS
0.16 (0.28)
0.05 (0.09)
0.05 (0.10)
0.15 (0.26)
0.62 (0.77)
– –
Note: The regions are designated as in Table 1. The highest values of similarity coefficients are shown in bold.
position by the absence of representatives of Gymnoscelis and Chloroclystis, and also by the presence of species from the genera Nasusina and Prorella (the similarity coefficients being 0.17 and 0.29). It shows the greatest similarity with the fauna of the Neotropical Region (0.20 and 0.33, respectively). The zoogeographic analysis is complicated by synonymy, which is typical of taxonomically difficult groups such as the tribe Eupitheciini. Based on the data on different geometrid subfamilies and the faunas of different biogeographic regions, Gaston and coauthors (1995) noted the direct proportion between the total number of species and that of synonymized names; however, such dependence is true only for the pug fauna of the Palaearctic (486 valid and 388 synonymized names) and has not been confirmed so far ENTOMOLOGICAL REVIEW Vol. 92 No. 4 2012
for other biogeographic regions. The strongest deviations from this trend can be observed for the faunas of the Nearctic and Neotropical regions (Fig. 3). In the first case, there are too many synonyms for such a small number of species; in the second, they are too few for considerable species diversity. The number of synonymized names will certainly grow after revision of Eupitheciini from the Afrotropical, Neotropical, Oriental, and Australian regions. CONCLUSIONS (1) The tribe Eupitheciini dominates by the number of species in the family Geometridae. The tribe unites almost 1900 species of the world fauna (about 3000 described species-group taxa) from 47 genera, 15 of which (almost a third) are monotypic.
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(2) The greatest diversity of pugs at the genus level is observed in the Australian (38 genera of which 11 are endemic) and Oriental (32 genera, 4 endemic) regions. In the Neotropical Region they are represented by only one genus. By the species richness, the biogeographic regions can be arranged in a different order: Palaearctic (487 species), Oriental (397), Neotropical (346), Australian (251), Afrotropical (198), and Nearctic (166). (3) The largest genus Eupithecia is the most abundantly represented in the Palaearctic (466 species) where it comprises more than 95% of the tribe Eupitheciini. Species of this genus are also well represented in the continental part of the Oriental Region. However, the number of Eupithecia species is abruptly reduced toward the equator, so that on the Sunda Islands it already comprises only about 2% of the tribe fauna. The fauna of the Australian Region includes only about 20 species of this genus, or about 8% of all the pugs. (4) The greatest similarity at the genus level is observed between the faunas of the Oriental and the Australian regions (the Jaccard and Sørensen coefficients being 0.62 and 0.77, respectively). The Palaearctic fauna is closer to the Afrotropical and Oriental than to the Nearctic one. The faunas of the Nearctic and Neotropical Regions are characterized by greater similarity (0.20 and 0.33) to each other than to the faunas of other biogeographic regions. (5) The greatest fraction of synonymized speciesgroup taxa is characteristic of the most fully studied faunas of Europe (64%) and North America (39%). On the contrary, the lowest level of synonymy can be observed in little studied, still unrevised faunas of the Afrotropical (8%) and Neotropical (9%) regions. ACKNOWLEDGMENTS The work was financially supported by the Ministry of Education and Science of the Russian Federation, the Russian Foundation of Basic Research (grant no. 11-04-01119), DFG (no. 436 RUS 17/99/02 and no. STU 498/2-1), and by a grant from the British Royal Society (2006). For the provided possibility to study the collections and for extensive help, the author is grateful to Dr. M. Scoble and Sir A.C. Galsworthy (Natural History Museum, London), Dr. D. Stüning (Zoological Research Institute and A. Kӧnig Museum, Bonn), Dr. A. Hausmann (Zoological Museum of Bavaria, Munich), Dr. W. Mey (Zoological Museum of
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