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Leaf-mining Nepticulidae (Lepidoptera) from record high altitudes: Documenting an entire new fauna in the Andean páramo and puna

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The monograph treats 29 species of leaf-mining pygmy moths (Insecta, Lepidoptera, Nepticulidae) discovered in the northern Andean bush and grass páramo and the central Andean puna at altitudes above 3700 m. They represent the world’s highest-altitudinal Nepticulidae fauna known. The height record belongs to Stigmella nivea sp. nov. from Peru collected at an elevation of 4700 m. Except for one species, all belong to Stigmella Schrank. Twenty-two of the species recorded are new and are named and described in the current paper: Stigmella paramica Diškus & Stonis, sp. nov.; S. lachemillae Diškus & Stonis, sp. nov.; S. gynoxyphaga Diškus & Stonis, sp. nov.; S. calceolariae Diškus & Stonis, sp. nov.; S. rigida Diškus & Stonis, sp. nov.; S. altiplanica Diškus & Stonis, sp. nov.; S. robusta Remeikis & Stonis, sp. nov.; S. pseudorobusta Remeikis & Stonis, sp. nov.; S. auriargentata Remeikis & Stonis, sp. nov.; S. altimontana Remeikis & Stonis, sp. nov.; S. pandora Remeikis & Stonis, sp. nov.; S. ampla Diškus & Stonis, sp. nov.; S. evanida Diškus & Stonis, sp. nov.; S. mustelina Remeikis & Stonis, sp. nov.; S. angusta Diškus & Stonis, sp. nov.; S. alticosma Remeikis & Stonis, sp. nov.; S. nivea Remeikis & Stonis, sp. nov.; S. kristenseni Diškus & Stonis, sp. nov.; S. lobata Remeikis & Stonis, sp. nov.; S. ageratinae Diškus & Stonis, sp. nov.; S. clinopodiella Diškus & Stonis, sp. nov.; and S. calceolarifoliae Diškus & Stonis, sp. nov. Some of these species are leaf-miners on Asteraceae (Pentacalia, Baccharis, Gynoxys, and Ageratinaplants), Calceolariaceae (Calceolaria), Lamiaceae (Clinopodium), and Rosaceae (Lachemilla). Twenty species are known only from adults with no data on their biology and host-plants. In addition, we present data and discuss recently discovered nepticulid taxa associated with Polylepis forests that is the natural vegetation in much of the High Andes. All High-Andean Stigmella species treated are illustrated with photographs of the adults and genitalia, distribution maps, including some with photographs of the leaf-mines and habitats.
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... In some of the literature Glaucolepis (=Fedalmia Beirne, 1945, synonymized by Puplesis 1985) has been recognized and presented as a separate, "good" genus (ie. not as a subgenus), e.g. by Wilkinson & Scoble (1979); Puplesis (1985Puplesis ( , 1992Puplesis ( , 1994; Puplesis et al. (1996); Puplesis & Robinson (2000); ; ; Navickaitė et al. (2011Navickaitė et al. ( , 2014a; Stonis et al. (2016Stonis et al. ( , 2017; Remeikis (2017). In some other publications Glaucolepis (=Fedalmia Beirne) was treated as a subgenus of Trifurcula Zeller, 1848 (e.g., Scoble 1983;van Nieukerken 1986a, b;Johansson et al. 1990;Laštůvka & Laštůvka, 2000;van Nieukerken et al. 2004;Laštůvka et al. 2007Laštůvka et al. , 2013Ivinskis et al. 2012). ...
... Currently, Glaucolepis comprises 42 named species worldwide: from the Americas, ie. the Western Hemisphere (Braun 1917;Wilkinson & Scoble 1979;Puplesis & Robinson 2000;Puplesis et al. 2002;Stonis et al. 2016Stonis et al. , 2017; Europe, reaching the highest diversity in the Mediterranean (e.g. Chrétien 1904Chrétien , 1907Chrétien , 1914Parenti 1963;Klimesh 1975Klimesh , 1978Klimesh , 1979Johansson et al. 1990; van Nieukerken & Puplesis 1991;Puplesis 1994;Laštůvka & Laštůvka 1997, 1998, 2007van Nieukerken et al. 2004Laštůvka et al. 2013;Navickaitė et al. 2014a, b); North Africa (Chrétien 1914(Chrétien , 1915; Asia (Puplesis 1985; van Nieukerken & Puplesis 1991;Puplesis 1994;Puplesis et al. 1996;, including one species from southwestern India ; and South America (Puplesis & Robinson 2000;Puplesis et al., 2002;van Nieukerken et al. 2016a;Stonis et al. 2016Stonis et al. , 2017, including two new species described from Argentina and Chile in this paper. ...
... Currently, Glaucolepis comprises 42 named species worldwide: from the Americas, ie. the Western Hemisphere (Braun 1917;Wilkinson & Scoble 1979;Puplesis & Robinson 2000;Puplesis et al. 2002;Stonis et al. 2016Stonis et al. , 2017; Europe, reaching the highest diversity in the Mediterranean (e.g. Chrétien 1904Chrétien , 1907Chrétien , 1914Parenti 1963;Klimesh 1975Klimesh , 1978Klimesh , 1979Johansson et al. 1990; van Nieukerken & Puplesis 1991;Puplesis 1994;Laštůvka & Laštůvka 1997, 1998, 2007van Nieukerken et al. 2004Laštůvka et al. 2013;Navickaitė et al. 2014a, b); North Africa (Chrétien 1914(Chrétien , 1915; Asia (Puplesis 1985; van Nieukerken & Puplesis 1991;Puplesis 1994;Puplesis et al. 1996;, including one species from southwestern India ; and South America (Puplesis & Robinson 2000;Puplesis et al., 2002;van Nieukerken et al. 2016a;Stonis et al. 2016Stonis et al. , 2017, including two new species described from Argentina and Chile in this paper. Doorenweerd et al. (2016) stated that more new species could possibly occur in East Asia and other regions. ...
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We provide diagnostic characters for the genus Glaucolepis Braun, re-examine the type series of the type species of the North American G. saccharella Braun, describe two new species from Chile and Argentina (G. flagellata Remeikis & Stonis, sp. nov. and G. pseudoflagellata Remeikis & Stonis, sp. nov.), and provide the first photographic documentation of the central Andean G. aerifica (Meyrick). We synonymize Neotrifurcula van Nieukerken, 2016, syn. nov. with Glaucolepis and provide one new combination for the south Andean G. gielisorum (van Nieukerken, 2016), comb. nov. All species treated in the paper are illustrated with drawings and (or) photographs of the adults and genitalia.
... The family Nepticulidae is characterized in several monographic reviews, notably by Newton & Wilkinson (1982), Scoble (1983), Kemperman & Wilkinson (1985), van Nieukerken (1986), Johansson et al. (1990), Puplesis (1994), Puplesis & Robinson (2000), Puplesis & Diškus (2003), Diškus & Stonis (2012) and Stonis et al. (2016d). ...
... Remeikis et al. (2016) reported that Lamiaceae feeders might make up a significant portion of the South American fauna of Nepticulidae. However, until now, except for the central Andean Stigmella clinopodiella Diškus & Stonis feeding on Clinopodium axillare (Stonis et al. 2016d), no other published evidence of this evidently diverse fauna was provided. This paper is the first to describe the Lamiaceae-feeding Stigmella Schrank species from the northern Andes: Stigmella lamiacifoliae Remeikis & Stonis, sp. ...
... Methods and protocols for species identification and description are outlined in Puplesis (1994), Puplesis & Robinson (2000), Puplesis & Diškus (2003), Diškus & Stonis (2012), and Stonis et al. (2014aStonis et al. ( , 2016d. After maceration of the abdomen in 10% KOH and subsequent cleaning, male genital capsules were removed from the abdomen and mounted ventral side uppermost. ...
Article
We provide a report on Nepticulidae feeding on Lamiaceae plants in South America and describe two new species from the Andes: Stigmella lamiacifoliae Remeikis & Stonis, sp. nov., feeding on Salvia palifolia in Colombia, and S. scutellariae Remeikis & Stonis, sp. nov., feeding on Scutellaria volubilis in Ecuador. The leaf mines, adults, genitalia, and habitats of the new species are illustrated.
... They are a specialized but phylogenetically primitive Lepidoptera family which comprises the smallest moth on Earth. The family was extensively characterized in monographic reviews by Scoble (1983), Johansson et al. (1990), Puplesis (1994), and Puplesis, Diškus (2003), with special reference to South America also by Puplesis, Robinson (2000) and Stonis et al. (2016c). One of the most prominent characteristics of the Nepticulidae is their larval biology. ...
... The host plants of only a small proportion of the species were known and include host plants in the Fabaceae (Leguminosae), Malvaceae, Polygonaceae (see Puplesis, Robinson, 2000;Puplesis et al., 2002aPuplesis et al., , 2002b. The history of Neotropical and Ando-Patagonian Nepticulidae research is provided by Stonis et al (2016c). Recently a first review of Rosaceae-feeding Nepticulidae (Stonis et al., 2016b(Stonis et al., , 2016d) and a few other papers dealing with new host-plant data (Stonis et al., 2015(Stonis et al., , 2016a have focused on Nepticulidae. ...
... Male genitalia (Figs. [12][13][14][15][16]. Capsule a little longer (320-340 μm) than wide (190-200 μm). ...
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The paper presents the first documentation of Urticaceae-feeding Nepticulidae species in South America and describes four new species: two species feeding on Phenax Wedd. (Stigmella singularia Diškus & Stonis, sp. nov. and S. lata Diškus & Stonis, sp. nov.), one species on Boehmeria Jacq. (S. boehmeriphaga Diškus & Stonis, sp. nov.), and one species on Pilea Lindl. (S. auripurpurata Diškus & Stonis, sp. nov.); all from the equatorial Andes. In addition, leaf-mines of an unknown Stigmella taxa feeding on Phenax are documented. The newly discovered Urticaceae-feeding Nepticulidae exhibit some morphological and taxonomical diversity: two species groups, Stigmella singularia and S. marmorea, are revealed (the latter is designated in the current paper).
... Pygmy moths (Nepticulidae) are a peculiar, specialized family of primitive (monotrysian) leaf-mining Lepidoptera with worldwide distribution. For a long period of time, Central and South America, though vast and with a hugely diverse biota, was comparatively unexplored regarding the collection and study of pygmy moths (for the history of research see Stonis et al. 2016d). However, recent studies showed that Nepticulidae and particularly its genus Stigmella Schrank are widespread in this region and particularly in the Andes (see Puplesis & Robinson 2000;Puplesis et al. 2002aPuplesis et al. , 2002bStonis et al. 2014bStonis et al. , 2015aStonis et al. , 2015bStonis et al. , 2016aStonis et al. , 2016bStonis et al. , 2016cStonis et al. , 2016dRemeikis & Stonis 2015;. ...
... For a long period of time, Central and South America, though vast and with a hugely diverse biota, was comparatively unexplored regarding the collection and study of pygmy moths (for the history of research see Stonis et al. 2016d). However, recent studies showed that Nepticulidae and particularly its genus Stigmella Schrank are widespread in this region and particularly in the Andes (see Puplesis & Robinson 2000;Puplesis et al. 2002aPuplesis et al. , 2002bStonis et al. 2014bStonis et al. , 2015aStonis et al. , 2015bStonis et al. , 2016aStonis et al. , 2016bStonis et al. , 2016cStonis et al. , 2016dRemeikis & Stonis 2015;. Therefore, it was not unexpected that our recent study of a large collection sample from the central Andes deposited in the ZMUC and other collections, revealed a diverse fauna of Nepticulidae, particularly of Stigmella, in the Peruvian and Bolivian Andes. ...
... Distribution of the central Andean Nepticulidae. 1, high-Andean species recorded at altitudes above 3700 m (after Stonis et al. 2016d); 2, species recorded at altitudes below 3700 m; 3-8, some collecting areas in the Peruvian Andes. ...
Article
We review forty-five species of Nepticulidae (Lepidoptera) based on an analysis of samples collected in the central Andean region of Peru and Bolivia. Thirteen of these species are new to science, and are named and described here: Stigmella paracosma Remeikis & Stonis, sp. nov., S. expressa Remeikis & Stonis, sp. nov., S. acalyphae Diškus & Stonis, sp. nov., S. lepida Diškus & Stonis, sp. nov., S. misera Diškus & Stonis, sp. nov., S. inca Diškus & Stonis, sp. nov., S. eiffeli Diškus & Stonis, sp. nov., S. arequipica Remeikis & Stonis, sp. nov., S. coronaria Diškus & Stonis, sp. nov., S. azulella Diškus & Stonis, sp. nov., S. sparsella Diškus & Stonis, sp. nov., Manoneura forcipis Diškus & Stonis, sp. nov., and Acalyptris murex Diškus & Stonis, sp. nov. Some of the central Andean species are recorded here as leaf-miners on Euphorbiaceae (Acalypha), Fabaceae (Collaea), Rosaceae (Polylepis), Malvaceae (Sida), Calceolariaceae (Calceolaria), Lamiaceae (Clinopodium), and Asteraceae (Ageratina and Trixis). We create eleven new species groups based on morphological characters designated in Stigmella and one in Acalyptris. A pictorial key to the species groups and distribution maps are provided. All new species are illustrated with 150 photographs and drawings of the adults and genitalia, and, where known and/or available, photographs of host-plants and leaf-mines.
... By comparing diversification rates of lineages in fast evolving biomes and by using numerous molecular phylogenies, it has been found that average diversification rates of páramo lineages are faster than those of other reportedly fast evolving biodiversity hotspots (Madriñán et al. 2013). In the páramo, organisms that populate this ecosystem are a likely product of specific adaptations to an extreme environment that evolved during the Pleistocene or the last three to five million years (Madriñán et al. 2013, Stonis et al. 2016. Thus, according to Madriñán et al. (2013), the páramo represents an ideal model ecosystem for investigating diversification processes. ...
... Geographical distribution of the genus Brachinepticula Diškus & Stonis. 48, distribution map of all currently known species; 49, South American transition zone of the Neotropical Region (afterStonis et al. 2016and Morrone 2014, 2015 ...
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This paper describes two new species of Brachinepticula Stonis & Diškus: B. melania Remeikis, Mey & Stonis, sp. nov. and B. colombica Remeikis, Mey & Stonis, sp. nov. Both the new species were collected in the Northern Andean Páramo (Cundinamarca, Colombia). Since the specimens were caught at night-time with a light trap, the host plants remain unknown. The examination of the morphology of the male genitalia of B. melania and female genitalia of B. colombica revealed a highly distinctive new taxa of pygmy moths and broadened our knowledge of the morphology and distribution of the recently described endemic genus Brachinepticula Diškus & Stonis. The examination of the new findings also allowed us updating the diagnosis of Brachinepticula. The new species were illustrated with photographs of the genitalia, adults, and habitats.
... In recent decades, the study of these particular lepidopteran families has been intensive and productive as never before. Therefore, a global catalog published in 2016 ( van Nieukerken et al. 2016a) which reviews Nepticulidae and Opostegidae, does not include approximately 14% of recently described species of the global Nepticulidae, primarily from Central and South America (species were listed in the following papers by Stonis et al. 2016cStonis et al. , 2017aStonis et al. , 2017cStonis et al. , 2018bStonis et al. , 2018cStonis et al. , 2018dStonis et al. , 2020cDobrynina et al. 2021). ...
... Nepticulidae, or pygmy moths (Figs 1-14), of Early Cretaceous origin (Doorenweerd et al. 2016), are distributed worldwide and occur in most terrestrial ecosystems, from tundra and temperate forests to arid boreal or subtropical deserts and various tropical habitats, including the lowland rainforest up to the elevations 4,700 m in the northern Andean bush and grass páramo and the central Andean puna (Stonis et al. 2016c). Externally, pygmy moths are characterized by an expanded scape of the antenna or eye cap, a distinctive frontal tuft of piliform scales on the vertex of the head, a lamellar (or piliform) collar on the back of the head, and a simple wing pattern comprised of one (or two) fascia(e) or spots, or with uniformly coloured forewings. ...
Article
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This study identifies the number of named and described species of three monotrysian, plant-mining lepidopteran families worldwide: Nepticulidae and Opostegidae (Nepticuloidea), and Tischeriidae (Tischerioidea). At the end of 2021, we estimated that a total of 1000 Nepticulidae species, 197 Opostegidae species, and 170 Tischeriidae species have been described since the taxonomic practice of describing species began in the 18th century. We examine and discuss the history of descriptions and authorship of species worldwide for each of the three families. We found that the total (accumulative) number of species described increased with each time period delineated. About five new species were described per year on average, or about 22 new species were described per year in the 21st century. We recognize researchers with the most number of described species in these three taxa.
... Nepticulid moths are a phylogenetically basal lepidopteran family and are famous for being the world's smallest, with a wingspan often less than 4-5 mm. Useful and extensive characterizations of this family were provided in several monographic reviews, notably those by Newton and Wilkinson (1982), Scoble (1983), Kemperman and Wilkinson (1985), van Nieukerken (1986), Johansson et al. (1990), Puplesis (1994), Puplesis and Robinson (2000), Puplesis and Di skus (2003), Di skus and Stonis (2012), and Stonis et al. (2016e). ...
... Methods and protocols for species identification and description are outlined in Puplesis (1994), Puplesis and Robinson (2000), Puplesis and Di skus (2003), and Stonis et al. (2016e). ...
Article
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Pygmy moths (Nepticulidae) associated with Asteraceae are poorly known and very rare worldwide. Recently, we discovered many leaf-mining nepticulids in South America feeding on Asteraceae. We review all known records of Asteraceae-feeding Nepticulidae, which in the Neotropics (including the Andes and Patagonia) previously included only Stigmella Schrank. We describe six new species of Stigmella from equatorial South America: S. jungiae Diškus and Stonis, n. sp. (feeding on Jungia L.f.), S. aeneola Diškus and Stonis, n. sp., S. violea Diškus and Stonis, n. sp., S. bracteata Diškus and Stonis, n. sp. (feeding on Liabum Adans.), S. spatiosa Diškus and Stonis, n. sp. (feeding on Ageratina Spach), and S. auripennata Diškus and Stonis, n. sp. (feeding on Baccharis L.). All new taxa are illustrated with photographs of the adults, their genitalia, and leaf mines. Additionally, leaf mines of three unknown nepticulid taxa on Jungia (aff. J. polita Griseb.), Piptocoma discolor (Kunth) Pruski, Gynoxys laurifolia (Kunth) Cass., and G. acostae Cuatrec. are documented for the first time. We diagnose and designate two new species groups and one new species complex in Stigmella. We discuss the origin and diversity of Asteraceae in the Neotropics and illustrate the distribution of Stigmella species feeding on representatives of Asteraceae at the tribal level.
... Male genitalia (Figs. 2,(12)(13)(14)(15)(16)(17)(18)(19)(20)(21)(22)(23)(24)(25)(26)(27)(28)(29)(30). Capsule longer (285-380 μm) than wide (140-280 μm). ...
... Phallus angular (Figs. [28][29][30] Female genitalia (Fig. 32). Relatively short, total length about 490 μm. ...
Article
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We describe one new genus (Brachinepticula Stonis & Diškus, gen. nov.) with two new species (B. plurilobata Diškus & Stonis, sp. nov., B. elongata Remeikis & Stonis, sp. nov.), and one species with uncertain taxonomic position (Johanssoniella bina Remeikis & Stonis, sp. nov.). We also provide diagnostic characters and an updated annotated catalogue of the following related genera: newly restored Johanssoniella Koçak, Brachinepticula gen. nov., Enteucha Meyrick (s. str.) and Manoneura Davis. In the Annotated Catalogue, we provide five new combinations and new data on morphology, biology, and distribution of some species, including the first photographic documentation of Manoneura basidactyla Davis discovered in the Amazon rainforest, results of re-examination of the male genitalia of the European Johanssonia acetosae (Stt.), also the first documentation of the male genitalia, host plant and leaf mines of the little known Johanssoniella diplocosma (Meyrick) from the Himalayas. All new taxa treated in the paper are illustrated with drawings and (or) photographs of the adults and genitalia; photographs of the leaf mines of Brachinepticula plurilobata Diškus & Stonis, sp. nov. and Johanssoniella diplocosma (Meyrick) are also provided.
... The family Nepticulidae (pygmy moths) has been characterized in monographic reviews by Johansson et al. (1990) and Puplesis (1994) and, with special reference to South America, by Puplesis, Robinson (2000) and Stonis et al. (2016). ...
... The description of the new species is based on material deposited in the collection of the Zoological Museum, Natural History Museum of Denmark in Copenhagen, Denmark, which was collected in Argentina in 1981 by E. S. Nielsen and Ole Karsholt during the international expedition Mision Cientifica Danesa. Collecting methods and protocols for species identification and description are outlined in Puplesis, Robinson (2000) and Stonis et al. (2016). After maceration of the abdomen in 10% KOH and subsequent cleaning, male genital capsules were removed from the abdomen and mounted ventral side uppermost. ...
Article
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The paper deals with a description of a new leaf-mining Nepticulidae species, Stigmella lilliputica Remeikis & Stonis, sp. nov., from Argentina, the larvae of which feed on leaves of the Chilean peppertree Schinus polygama (Cav.) Cabrera. The species is illustrated with photographs of the adults and leaf mines, a distribution map as well as drawings and photographs of the genitalia.
... The family was characterized with special reference to Middle and South America in monographic reviews by Puplesis, Robinson (2000) and Stonis et al. (2016b); however, very little information on host-plants and other particularities of the biology of Nepticulidae was provided. ...
... Larvae of Nepticulidae (pygmy moths) are miners during all instars and feed inside leaves and occasionally other assimilation tissues of plants (Figs. [1][2][3][4][8][9][10]. One of the most pronounced and well-known trophic particularities is narrow diet breadth (stenophagy): most of Nepticulidae are either monophagous or strict oligophagous and rarely broad oligophagous, and only in a few known cases disjunct oligophagous (for the definition of the terms and the predominance of monophagy in Nepticulidae, see Diškus, Stonis, 2012 andPuplesis, Diškus, 2003). ...
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The paper briefly analyzes trophic relationships of Nepticulidae occurring in Middle and South America (the Neotropical and Ando-Patagonian regions) for the first time. The results show some peculiar features, including unexpectedly large percentage of taxa trophically associated with Asterales (Asteraceae) and Lamiales plants. However, because of insufficient sampling in most regions of the Caribbean and Middle and South America as well as the fact that only 50% of analyzed taxa are known in host-plant records, the results of current analysis should be treated as preliminary.
... During recent years, the inventory of the Neotropical fauna of Nepticulidae was very active (e.g. Puplesis & Robisnon 2000;Puplesis et al. 2002aPuplesis et al. , 2002bStonis et al. 2016Stonis et al. , 2017Stonis et al. , 2019Stonis et al. , 2020Stonis et al. , 2021Remeikis & Stonis 2015;van Nieukerken et al. 2016;Stonis & Remeikis 2018). The description of the overlooked species, O. rionegrella sp. ...
Article
We describe a new species, Ozadelpha rionegrella Remeikis & Dobrynina, sp. nov., a pygmy moth (Nepticulidae) from the southern Andes (Argentina). The new species is illustrated with photographs of the adult and male genitalia. We briefly discuss the issues of diagnostics of the new species and the related O. ovata (Puplesis & Robinson, 2000) and provide an updated pictorial tool for diagnostics of all currently known Ozadelpha species.
... A few of them are listed further in the text, and a small, freely selected portion of further uncited references are provided here as principally important papers for the current study: Wilkinson (1979); Wilkinson & Scoble (1979); Newton & Wilkinson (1982); Scoble (1983); Kemperman & Wilkinson (1985); Donner & Wilkinson (1989); van Nieukerken (1985avan Nieukerken ( , 2007avan Nieukerken ( , 2007bvan Nieukerken ( , 2019; Hoare et al. (1997); Hoare (2000aHoare ( , 2000b; Puplesis & Robinson (2000); van Nieukerken & Liu (2000); ; ; ; and Hoare & van Nieukerken (2013). Recently, based on new discoveries of Nepticulidae in the Neotropics, many new measurement data became available because Stonis et al. (2013aStonis et al. ( , 2013bStonis et al. ( , 2016Stonis et al. ( , 2017aStonis et al. ( , 2017bStonis et al. ( , 2017cStonis et al. ( , 2018aStonis et al. ( , 2018bStonis et al. ( , 2018cStonis et al. ( , 2020b; ; , and many others (a few are cited further in the text). ...
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The world’s smallest moths in Lepidoptera (Insecta) and the complexity in making such a determination are examined and discussed. The forewing length and wingspan of 650 species were measured and the same data were retrieved from published papers to determine which species and family have the smallest moths in the world. The minimal recorded forewing length was found to be around 1.2–1.3 mm and the wingspan around 2.6–2.8 mm in two families, the Gracillariidae and Nepticulidae. Among Lepidoptera, the following species have the smallest moths globally: the European Johanssoniella acetosae (Stainton), the Peruvian Simplimorpha kailai Stonis & Diškus, the Mexican Stigmella maya Remeikis & Stonis, the Mediterranean S. diniensis (Klimesh), the Mediterranean Parafomoria liguricella (Klimesh) (Nepticulidae), the South East Asian Porphyrosela alternata Kumata, and the Central African P. desmodivora De Prins (Gracillariidae). Additionally, in the Nepticulidae, we provide a measurement update for Stigmella maya Remeikis & Stonis, one of the tiniest species with a forewing length of 1.3 mm and wingspan of 2.8 mm, and describe a new species, Stigmella incaica Diškus & Stonis, sp. nov., with a forewing length of 1.75 to 1.95 mm and a wingspan of 3.8 to 4.3 mm.
... Collecting methods, including rearing of adults from mining larvae, were outlined in Puplesis (1994), Diškus & Stonis (2012), and Stonis et al. (2016). Protocols for species identification and description were outlined in Johansson et al. (1990), Puplesis (1994), and Puplesis and Diškus (2003). ...
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We review eighteen species of Nepticulidae (Lepidoptera: Nepticuloidea) from Asia and describe eleven new species from Central Asia and the western Himalaya: Stigmella ziziphifolia Rocienė & Stonis, sp. nov., S. damocles Remeikis, sp. nov., S. pyramidata Diškus & Navickaitė, sp. nov., S. alilediella Diškus & Navickaitė, sp. nov., S. longa Remeikis & Stonis, sp. nov., S. latilobata Diškus & Navickaitė, sp. nov., S. paniculata Diškus & Navickaitė, sp. nov., Ectoedemia orbiculata Diškus, Remeikis & Stonis, sp. nov., Acalyptris brunipexus Stonis, Diškus & Remeikis, sp. nov., A. noctilucus Rocienė & Stonis, sp. nov., and A. nasutus Diškus & Navickaitė, sp. nov. The new taxa are illustrated with photographs of the adults, their genitalia, and, if available, leaf mines. We also provide the first photographic documentation of adults and male genitalia of some Asiatic species. We synonymize E. petrosa Puplesis with E. spinosella (de Joannis), and E. rosiphila Puplesis with E. ingloria Puplesis.
... The history of documentation of the Nepticulidae fauna of the Neotropics counts about 133 years with the description of the first species from Colombia (see Puplesis, Robinson, 2000), but only during the recent decade it has become really active. A series of taxonomic publications on Nepticulidae covering Central and South America were published recently, notably, Remeikis et al., 2014;Stonis et al., 2013Stonis et al., , 2014bStonis et al., , 2016aStonis et al., , 2016bStonis et al., , 2016cStonis et al., , 2016dStonis et al., , 2017bStonis et al., , 2017cStonis et al., , 2018aStonis et al., , 2018bStonis et al., , 2018cStonis et al., , 2018dStonis et al., , 2019van Nieukerken et al., 2016avan Nieukerken et al., , 2016bStonis, Diškus, 2018, etc. Here, we continue to document the Nep­ ticulidae fauna of the Neotropics and de­ scribe one more species, Ozadelpha nigella Diškus & Stonis, sp. ...
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We describe a new species, Ozadelpha nigella Diškus & Stonis, sp. nov., a leaf miner on Symplocos Jacq. (Ericales: Symplocaceae), from the Yungas biogeographical province, South America. We briefly discuss the issues of diagnostics of the recently erected genus Ozadelpha van Nieukerken. The genus is now comprised of four named species having shared characters useful for efficient identification and genus recognition. The new species is illustrated with photographs of the adults, male genitalia and leaf mines.
... We followed collecting methods and protocols for species description outlined in Johansson et al. (1990), Puplesis and Diškus (2003), and Stonis et al. (2016). After maceration of the abdomen in 10% KOH and subsequent cleaning, male genital capsules were removed from the abdomen and mounted ventral side up. ...
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We describe a new pest of guava ( Psidium guajava L.), Hesperolyra guajavifoliae Stonis & Vargas, sp. nov. , that was recently discovered in western Colombia. Hesperolyra van Nieukerken is a small, Neotropical genus of pygmy moths (Nepticulidae). We re-examine and document the complex morphology of the male genitalia of the generic type species, H. diskusi (Puplesis & Robinson). We discuss the diagnostics and composition of the genus and provide a simple pictorial differentiation scheme for all currently known representatives of the genus. The new species is illustrated with photographs of the adults, some of the immature stages, male and female genitalia, and leaf mines. A link to the COI barcodes of H. guajavifoliaesp. nov. is provided and the relationship of Hesperolyra to other genera is discussed.
... Methods and protocols for species identification and description are outlined in Puplesis (1994), Puplesis and Robinson (2000), Puplesis and Diškus (2003), and Stonis et al. (2016). ...
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We describe two new species of Nepticulidae (Lepidoptera) from the Western Hemisphere that feed on Fabaceae: Stigmella mimosae Diškus and Stonis, new species, on Mimosa albida Willd. Caesalpinioideae), and Acalyptris minimus Diškus and Stonis, new species, on Centrolobium ochroxylum Rose ex Rudd. (Faboideae). Centrolobium ochroxylum is an economically important tree in South America that is exported for timber. The new species are illustrated with photographs of the adults, genitalia, and the leaf mines.
... Species groups are fully informal units. For diagnostic purposes, species groups are widely in use, e.g., in the Nepticulidae (see Stonis et al. 2013Stonis et al. , 2016bStonis et al. , 2017bStonis et al. , 2018 and Elachistidae (see e.g. Kaila 2011, 2015. ...
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We review eleven Astrotischeria Puplesis & Diškus (Lepidoptera: Tischeriidae) species which possess a novel character for the Tischeriidae family: a highly modified valva of the male genitalia with one ventral and two dorsal lobes (or processes). The species are distributed in the Americas, including the USA, Caribbean (St. Thomas), Central America (Belize, Guatemala and Honduras), and South America (Ecuador, Bolivia, and Brazil). Species for which the biology has been studied are associated with host plants from Asteroidea of the Asteraceae family. The following seven species are described as new: Astrotischeria trilobata Diškus & Stonis, sp. nov., A. amazonica Diškus & Stonis, sp. nov., A. maya Diškus & Stonis, sp. nov., A. selvica Diškus, Carvalho-Filho & Stonis, sp. nov., A. casila Diškus & Stonis, sp. nov., A. onae Diškus & Stonis, sp. nov., and A. furcata Stonis & Diškus, sp. nov. A new informal species unit, the A. trilobata group, is designated for diagnostic purposes despite some doubts about monophyly of the group. Astrotischeria longeciliata (Frey & Boll) is synonymized here with the North American A. helianthi (Frey & Boll), a species not belonging to the A. trilobata group, syn. nov. For the first time, a method of rearing of adults from mining larvae, specifically adopted for Tischeriidae, is detailed. All species treated in the paper are illustrated with photographs or drawings of the adults, male genitalia, and, if available, the female genitalia, leaf mines and habitats. A distribution map for the species of the A. trilobata group and a scheme of the trophic relationships of the global Tischeriidae fauna are also provided.
... The descriptive terminology of morphological structures mostly follows Johansson et al. (1990), Puplesis (1994), Puplesis & Robinson (2000), and Stonis et al. (2016). ...
Article
In addition to numerous new species that can be placed to genera, our recent study of a large collection sample of Nepticulidae (Lepidoptera) from Central and South America revealed a few odd-looking new species, the taxonomic position of which seems rather problematic and, therefore, preliminary. Here we describe three new species of pygmy moths (Nepticulidae) from the Amazonian rainforest (Venezuela) and southern Andes (Chile and Argentina) possessing uncommon morphology. We also provide the first photographic documentation of the Central American Acalyptris argentosa (Puplesis & Robinson, 2000) with rather odd and hitherto unknown hindwing scaling. All species treated in the paper are extensively illustrated with drawings and (or) photographs of the adults and genitalia.
... The pygmy leafmining moths or Nepticulidae have around 300 species in Europe, the great majority of which are leafminers, feeding on woody plants and they are abundant in forested areas (van Nieukerken et al. 2016). Overall they do not form an important aspect of the diversity of high mountain faunas (but see Stonis et al. (2016) for a group of Stigmella Schrank, 1802 species occurring in the high Andes). In Europe few Nepticulidae are adapted to high altitudes, and apart from some common species with large altitudinal ranges, only a few species of Stigmella are alpine specialists, particularly a group of species feeding on herbaceous Rosaceae (Klimesch 1981;Johansson and Nielsen 1990;Laštůvka and Laštůvka 1997). ...
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Stigmella tatrica sp. n. is described from moths taken in the alpine zone of the Tatra Mountains in Slovakia. The new species is similar to several other montane species of Stigmella Schrank, 1802 in the S. aurella (Fabricius, 1775) group in external characters and male genitalia; its closest relative is S. dryadella (O. Hofmann, 1868). It is indistinguishable from S. tormentillella (Herrich-Schäffer, 1860) by the colour and pattern elements of the forewing; however, it differs in the male genitalia in the shape and number of cornuti, in the female genitalia by long apophyses with protruding ovipositor, and by COI barcodes. Immature stages are unknown, but Dryas octopetala L. (Rosaceae) is the possible hostplant. The long ovipositor suggests an unusual, possibly hidden place for oviposition. The male and female adults and genitalia of both sexes are figured and photographs of the habitat are provided.
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For the first time, genera of leaf mining Tischeriidae of the global fauna are reviewed and four new genera are described: Coptotrichoides Diškus & Stonis, gen. nov., trophically associated with Sapindaceae from Central and South America; Rytietia Diškus, Xu & Dai, gen. nov., trophically associated with Annonaceae from East and South East Asia; Pafazaria Diškus & Stonis, gen. nov., trophically associated with Fabaceae and Malvaceae from East and South East Asia; and Gnathitischeria Diškus, gen. nov., trophically associated with Malvaceae and Asteraceae from Central America. In addition to the descriptions of new genera, all previously described genera (Coptotriche Walsingham, 1890; Dishkeya Stonis, 2020; Tischeria Zeller, 1839; Manitischeria Diškus & Stonis, 2021; Neotischeria Diškus & Stonis, 2021; Paratischeria Diškus & Stonis, 2017; and Astrotischeria Puplesis & Diškus, 2003) are characterized and a pictorial identification key for all 11 genera is provided. Morphological and biological characteristics of all genera are supplemented with some molecular data. We described 12 new species: South East Asian Coptotriche sapaensis Diškus & Stonis, sp. nov.; Central American Coptotrichoides sapindoidum Diškus & Stonis, sp. nov. and Tischeria neokristenseni Diškus & Stonis, sp. nov.; South East Asian Rytietia uncinata Diškus, Xu & Dai, sp. nov., East Asian R. chongyiensis Xu & Dai, sp. nov.; South East Asian Pafazaria capitata Diškus & Stonis, sp. nov. and P. faboidica Diškus & Stonis, sp. nov.; Central American Neotischeria antigua Diškus & Remeikis, sp. nov. and N. subantigua Diškus & Remeikis, sp. nov.; South American Astrotischeria incae Diškus & Stonis, sp. nov.; and Central American Gnathitischeria atitlani Diškus & Stonis, sp. nov. and G. arcana Diškus & Stonis, sp. nov. The taxonomic rank of the East Asian subspecies Tischeria decidua siorkionla Kozlov, 1986 is elevated to species rank: T. siorkionla Kozlov (stat. nov.). The male genitalia of holotypes, paratypes, lectotypes or paralectotypes, or non-type specimens of some little-known species are examined for the first time. We provide the first morphological documentation for the following: Tischeria ceanothi lectotype and paralectotype; T. immaculata (stat. restored) paratype; Coptotrichoides deliquescens (Meyrick) lectotype and paralectotype; lectotypes of Coptotriche aenea (Frey & Boll), Astrotischeria plagifera (Meyrick), A. helianthi (Frey & Boll), and A. ephaptis (Meyrick); Coptotriche berberella (De Prins) paratype; and the first photographic documentation of Tischeria lvoskyi Kozlov holotype. Some non-type specimens deposited at NHMUK, including a few previously neglected species, are also examined and documented, including the American Astrotischeria solidagonifoliella (Clemens) and A. omissa (Braun). Nine new combinations are provided: Coptotrichoides deliquescens (Meyrick, 1915), comb. nov.; C. singularis (Stonis & Diškus) comb. nov.; C. suprafasciata (Diškus & Stonis) comb. nov., C. serjaniphaga (Remeikis & Stonis) comb. nov.; C. braziliensis (Diškus & Stonis) comb. nov.; Pafazaria jingdongensis (Xu & Dai), comb. nov.; Neotischeria explosa (Braun) comb. nov.; N. pallidipennella (Braun) comb. nov.; and Astrotischeria heteroterae (Frey & Boll) comb. nov. The synonymization of Tischeria longeciliata Frey & Boll, 1878 with Astrotischeria helianthi (Frey & Boll, 1878) is confirmed. One new species group, the Tischeria ceanothi group, is established. We also report a new distribution record of T. dodonaea Stainton from the Caucasus, Armenia. Three tables and 569 figures of adult external morphology, male and female genitalia, leaf mines, and cladograms based on molecular characters are provided.
Chapter
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It includes three separate chapters: Collecting of Nepticulidae Adults (by J. R. Stonis, A. Remeikis), Rearing of Nepticulidae Adults from Mining Larvae (by A. Diškus, J. R. Stonis), and Specimen Documentation and Micro-Mounts of Genitalia Structures Adopted for the Minute Lepidoptera (by J. R. Stonis, A. Diškus, A. Remeikis).
Book
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The publication deals with 228 species of Nepticulidae (or pygmy moths) from the Neotropics (i.e., the Neotropical and Ando-Patagonian regions) and adjacent areas. The species are illustrated with photographs or drawings of adult forewings, the male genitalia and, if available, leaf mines. In addition to the illustrated species review, the monograph announces about the launching of the first electronic diagnostic system (a free electronic tool on the internet) designed for the identification of species of Nepticulidae and includes chapters on the global biodiversity inventory, the collecting of adults and larvae of pygmy moths, preparation of micro-mounts of genitalia structures of Nepticulidae, global generic composition of the family, some particularities of Nepticulidae biology, a unique morphology guide based on the forewing pattern and male genitalia, descriptions of four newly named species, and a taxonomic checklist of the species currently known from the Neotropics and adjacent areas.
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Members of the Lamiaceae, or mint family, are used worldwide for medicinal, culinary and/or magical-religious purposes, as well as in pesticides and as ornamental plants. Very little is known about nepticulids, or pygmy moths, as leaf miners of Lamiaceae, but they may be an important component of South American diversity and potential pests of economically-important species of the mint family. In this paper, four new species of leaf-mining Nepticulidae are described from the equatorial Andes of Ecuador: S. mentholica Diškus & Stonis, sp. nov., Stigmella aromatica Diškus & Stonis, sp. nov., S. odora Diškus & Stonis, sp. nov., feeding on Minthostachys mollis (Benth.) Griseb., and S. tomentosella Diškus & Stonis, sp. nov., feeding on Clinopodium tomentosum (Kunth) Govaerts. It is hypothesized that host-plant distribution ranges can provide clues to potential distribution ranges of these newly discovered, trophically specialized leaf miners. The leaf mines, adults, and the genitalia of the new species are illustrated with photographs.
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We describe one new species (Acalyptris amazonensis sp. nov.) from the Amazonian rainforest of Ecuador and provide the first photographic documentation of two other related species also occurring in Ecuador (A. onorei Puplesis & Diškus and A. ecuadoriana Puplesis & Diškus). We briefly discuss diagnostic characters of the A. latipennata species group, which has an outstanding but disputable position within the genus Acalyptris Meyrick.
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The genus Simplimorpha Scoble is recognized for the first time from South America. We describe a subgenus, Myrtinepticula Stonis & Diškus, subgen. nov., for three new species from the southern Andes (Chile and Argentina): Simplimorpha (Myrtinepticula) cercaria Diškus & Stonis, sp. nov., S. (M.) nielseni Remeikis & Stonis, sp. nov., S. (M.) sapphirella Remeikis & Stonis, sp. nov.; and one new species from the southwestern Amazon (Peru): S. (M.) kailai Stonis, sp. nov. We provide a pictorial differentiation scheme for Simplimorpha Scoble and Pectinivalva Scoble. We synonymize the recently erected, predominantly Australian Menurella Hoare, syn. nov. and Cosanovula Hoare, syn. nov. with Pectinivalva Scoble. We also revise the taxonomic status of the Australian Roscidotoga Hoare as a subgenus of Simplimorpha which now exhibits a Gondwanan distribution in the Southern Hemisphere, with the presence of a single species in the Mediterranean. All new taxa are illustrated with photographs of the adults and the genitalia; the leaf mines of Simplimorpha (Myrtinepticula) nielseni sp. nov. are also provided.
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The paper reviews the most recent findings of the Rosaceae-feeding Nepticulidae species along with previous records of these tropically specialized leaf-miners in South America and describes three new species: one species on Hesperomeles obtusifolia (Pers.) Lindl (Stigmella circinata Diškus & Stonis, sp. nov.) and two species on Rubus spp. (S. rubiphagiella Diškus & Stonis, sp. nov. and Ectoedemia morae Diškus & Stonis, sp. nov.); all from the equatorial Andes. Additionally, leaf-mines of the unknown Stigmella taxa feeding on Acaena L., Alchemilla L., Rubus L., Prunus L., and Hesperomeles Lindl are documented. Description of previously unknown females of Stigmella nubimontana Puplesis & Diškus and photographic documentation of leaf-mines of S. nubimontana and S. rubeta Puplesis & Diškus are provided for the first time. The discovered Rosaceae-feeding Nepticulidae exhibit morphological and taxonomical diversity: two new species groups, Stigmella imperatoria and S. circinata, are designated; the latter is also compared with the most similar and probably closely related Holarctic S. hemargyrella and S. sorbi groups.
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In animals, adaptive radiation at the level of species is based to a large extent upon adaptations to survival in different climates, as well as to the occupation of different ecological niches in similar communities. In plants, adaptive radiation at the species level, as well as the origin of higher categories, depends to a large extent upon diversification of the methods of reproduction, both pollination and seed dispersal, as well as seed germination and the establishment of seedlings. With respect to the latter, evolutionary success depends to a large extent upon solution of three often conflicting problems: protection of the developing seed, dispersal to a new but similar location; and germination with sufficient seedling vigor to survive competition. In plants, the origin of higher categories is not associated with the appearance of distinctive characters that never serve to distinguish species. Evolutionary progress, here regarded as an increase in the amount and diversity of DNA-coded information, is based upon the bias in favor of mutations that increase rather than reduce the amount of genetic information already present.