Differentiation of Paratischeria and Neotischeria gen. nov. (Lepidoptera, Tischeriidae), with a description of new, mostly Asteraceae-feeding species from Central and South America

INTRODUCTION

Tischeriidae, or trumpet moths, represent a  relatively small family of leaf-mining lepidopterans. We expect over 300 species worldwide, but only about 175 Tischeriidae species have been documented so far. Many new species have already been collected and are ready for taxonomic description, but the  majority are still awaiting discovery, particularly from the  areas of South East Asia, equatorial Africa, and tropical America that are still poorly surveyed (Stonis, Solis, 2020). The  adults of trumpet moths possess numerous specialised morphological characters, including long antennal sensillae trichodea on the male antenna, a modified phallus in the male genitalia, or ovipositor lobes covered with thickened peg-like setae along with unique, rod-like or plate-like projections known as prela in the female genitalia. The  unique characters of Tischeriidae were overviewed in the monographs by Braun (1972) and Puplesis, Diškus (2003), as well as discussed and variously illustrated in recent papers, including those by Xu et al. (2017, 2018), Stonis et al. (2017, 2018a, 2019a, 2020a, 2020b, 2020c, 2021a, 2021b), and Stonis, Solis (2020).

Initially, the family comprised a single genus, Tischeria Zeller. However, before our study, Tischeridae was already comprised of six genera: Tischeria Zeller, 1839; Coptotriche Walsingham, 1890; Astrotischeria Puplesis & Diškus, 2003; Paratischeria Diškus & Stonis, 2017 (Stonis  et  al., 2017); Dishkeya Stonis, 2020 (Stonis, Solis, 2020), and Manitischeria Diškus & Stonis, 2021 (Stonis et al., 2021b).

In the  current paper, we briefly discuss the  diagnostics of Paratischeria, which is characterised by a  distinct dorsal sclerite in the male genitalia and the feeding of their larvae on Urticaceae, occasionally Malvaceae, host plants. Because of the absence of a dorsal sclerite, we exclude seven Asteraceae and Malvaceae-feeding species from Paratischeria and describe a new genus, Neotischeria Diškus & Stonis, gen. nov.

We also describe four new Neotischeria species discovered from Guatemala (Central America) and the  equatorial and central Andes (South America): N. conexa Diškus & Stonis, sp. nov., N. ochripennata Diškus & Remeikis, sp.  nov., N.  longa Diškus & Stonis, sp.  nov., and N.  poseidonia Diškus & Stonis, sp. nov. The host plant of the latter species remains unknown, and three other species are tropically associated with Liabum Adans and Smallanthus Mack. ex Small (Asteraceae).

Drawing upon our study of the  lectotype and paralectotype of Neotischeria capnota (Meyrick, 1915) deposited at NHMUK, we provide the first documentation of this little-known species.

MATERIALS AND METHODS

The documentation of taxa and description of the  new species is based on the  material deposited in the  collections of the  Zoological Institute of the  Russian Academy of Sciences, St. Petersburg, Russia (ZIN); some male and female paratypes of Neotischeria conexa sp. nov. and N. longa sp. nov. are also deposited at the  Nature Research Centre, Vilnius, Lithuania (NRC). Studies into the  lectotype and paralectotype of N.  capnota (Meyrick, 1915) are based on the material deposited at the Natural History Museum, London, United Kingdom (NHMUK).

Protocols regarding the  study into Tischeriidae have been previously described: the collecting and rearing of leaf mines by Stonis et al. (2018a), and techniques of specimen dissection and genitalia mounts in Euparal by Stonis et al. (2014). The descriptive terminology, with a  few exceptions, follows Puplesis, Diškus (2003).

The identifications of the Asteraceae host plants were undertaken by experts in the taxonomy of the family. Being based exclusively on field photographs (herbarium specimens were not collected and preserved) they should, therefore, be considered with caution.

Permanent preparations on microscope slides were photographed and studied with a  Leica DM2500 microscope and a  Leica DFC420 digital camera. Adults were measured and studied using Lomo stereoscopic microscopes MBS-10 and photographed using a  Leica S6D stereoscopic microscope with an attached Leica DFC290 digital camera.

For molecular analysis, genomic DNA was extracted from hind legs or whole adult specimen stored in 96% ethanol, using the Gene-Jet Genomic DNA Purification Kit (Thermo Fisher Scientific Baltics) following the manufacturer’s specifications. PCR reactions were carried out in an Eppendorf Mastercycler gradient 5331 (Germany). Cycling parameters were an initial denaturation step at 94°C for 2 min, followed by 94°C for 30 s, 50°C for 45 s, and 72°C for 1 min. This cycle was repeated 35 times, followed by 4 min of extension at 72°C. PCR products were visualized on 1.5% agarose gel stained with ethidium bromide and UV light photographs of the gels with DNA bands were taken using ‘Herolab’ transluminator (Germany). The  sequencing of some species was produced by the BigDye^® Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems) in Macrogen Inc. (Seoul, South Korea). The divergence was calculated using the  Kimura two-parameter distance (Kimura, 1980) model and graphically displayed in a Neighbour-Joining (NJ) tree by the  MEGA 7 software (Kumar et al., 2016). The robustness of the inferred tree was evaluated by bootstrapping with 10,000 replicates; distantly related Pseudopostega bogotensis Vargas (Opostegidae) was used as an outgroup. Molecular methods used in this paper for a fragment of the NJ tree (Fig.  1) are described in more detail in Stonis et al. (2019b).

DIFFERENTIATION OF PARATISCHERIA AND NEOTISCHERIA GEN. NOV.

The genus Paratischeria Diškus & Stonis was described for the species of specific morphology of the male genitalia (Stonis et al., 2017). This genus differs from Astrotischeria Puplesis & Diškus (described in Puplesis, Diškus, 2003) in the undivided, usually narrow valva (divided in Astrotischeria), long undivided uncus (usually short and divided in Astrotischeria), and the strongly, though variously, developed anellus (Stonis et al., 2017).

During current molecular studies into Tischeriidae species using fragments of the mitochondrial COI gene that included the standard barcode region and consequent studies on the obtained sequences of Paratischeria species, we found that there were two groups of the species which never clustered together in any of our phylogenetic trees but exhibited a clear dichotomy (Fig. 1).

The morphology of the  male genitalia also differed significantly: the  first group of the  species possessed only a  well-developed anellus (Figs 2–4), while the species of the second group were characterised by the presence of an anellus and a  distinct dorsal sclerite, a unique, derived character within Tischeriidae (Figs  5–7). The  feeding preferences also differed among the species of these two groups: the  host plants of the  first group belong to Asteraceae and Malvaceae, while the  species of the  second group feed on Urticaceae and occasionally on Malvaceae. By drawing on unpublished molecular data, the morphological characters that are easily applicable for further diagnostics, and differences in host-plant preferences, here we describe a new genus, Neotischeria gen. nov.

Because of the absence of a dorsal sclerite in the male genitalia, we exclude seven Asteraceae- and Malvaceae-feeding species from former Paratischeria, and provide seven new taxonomic combinations with Neotischeria: N. guarani (Diškus & Stonis, 2020) (comb. nov.), N. mesoamericana (Diškus & Stonis, 2020) (comb. nov.), N.  belizensis (Remeikis & Stonis, 2020) (comb. nov.), N.  robinsoni (Diškus & Stonis, 2020) (comb. nov.), N. tubifex (Diškus & Stonis, 2020) (comb. nov.), N.  neotropicana (Diškus & Stonis, 2015) (comb. nov.), and N.  capnota (Meyrick, 1915) (comb. nov.).

DESCRIPTION OF A NEW GENUS, WITH A REVIEW OF INCLUDED SPECIES

Neotischeria Diškus & Stonis, gen. nov.

urn:lsid:zoobank.org:act:14A682A5-F0B1-4352-A828-4C4AA150123D

Type species. Neotischeria neotropicana (Diškus & Stonis, 2015: 457–465).

Diagnosis. External characters or characters of wing venation are not informative or insufficient for generic differentiation in most cases of Tischeriidae, including this new genus. In the male genitalia, the combination of a slender, undivided, one-lobed valva, presence of a well-developed anellus, and the absence of a dorsal sclerite and transtilla distinguish the new genus from all other genera of Tischeriidae.

Description. Adult. Medium-small moths: forewing length 2.4–5.4  mm, wingspan 5.4–11.3 mm. Head: frons smoothly scaled; frontal tuft overlapping the  frons, comprised of long lamellar scales; collar comprised of slender lamellar scales; antenna longer than one half the length of forewing. Male forewing without a distinctive pattern, from bright yellow-ochre to pale grey cream, irregularly speckled with some darker scales; females often exhibit a remarkable sexual dimorphism and are characterised of a bright, patchy forewing; fringe line usually present. Hindwing slender, from cream or pale ochreous brown to dark grey-brown, without androconia. Male abdomen with anal tufts.

Male genitalia. Capsule significantly longer than wide. Uncus with two long or very long latero-ventral lobes and with (or without) small, rounded, dorsal lobes. Socii membranous, not thickened. Valva usually slender, occasionally widened in basal part, always without dorsal lobes, undivided (one-lobed); basal process long; transtilla absent. Juxta absent. Anellus well-developed, thickened, and with some setae laterally. Vinculum with a large or medium large, distally usually rounded or triangular ventral lobe. Phallus slightly or broadly bifurcated apically, without spines or carinae, except for N. robinsoni Diškus & Stonis and N. tubifex Diškus & Stonis, and N. mesoamericana Diškus & Stonis (those species possess carina-like processes on the phallus).

Female genitalia with large or medium-large rounded ovipositor lobes densely clothed with short, modified peg-like setae. Anterior and posterior apophyses usually almost equal in length; prela comprised of three pairs of rod-like projections. Corpus bursae with long or very long and very slender caudal part, and relatively small and slender main part; without pectinations or signum. Accessory sac absent or inconspicous. Ductus spermathaecae very slender, with many large or often very large coils.

Bionomics. Host plants are mainly from the Asteraceae family, including Liabum Adans., Smallanthus Mack., Montanoa Cerv., Elephantopus L., Otopappus Benth., Lasianthaea DC., but in the case of the N. neotropicana group, some host plants are from Malvaceae (Sida spp.). Larvae mine leaves and produce blotch-like leaf mines, without distorting the mined leaf (i.e., without bending the  leaf margin), except for N. neotropicana; the larva of the latter species distorts the mined leaf. Pupa in a circular, usually distinctive, nidus.

Distribution. Exclusively neotropical, endemic. Currently the  genus is known from subtropical Mexico to Paraguay and the Yungas biogeographical province of Bolivia, with the majority of species in the tropical habitats of Central America (Belize and Guatemala), occurring at an elevation from just few metres above sea level to 3000–3100 m in the equatorial Andes of South America.

Etymology. The  name Neotischeria (feminine) is a combination of the prefex neo (new), here derived from Neotropics where the genus is distributed, and the Latin name of another, iconic, type genus of the trumpet moth family, Tischeria.

Remarks. Unpublished molecular data provides support for this new genus. In our study, all sequences (with a final aligned length of the data-set 657 bp) have been deposited in the GenBank (www.ncbi.nlm.nih.gov/Genbank) and BOLD platform (www.barcodinglife.org); they will be published separately with a molecular analysis of all genera of Tischeriidae (Stonis et al., in prep.).

1. Neotischeria conexa Diškus & Stonis, sp. nov.

urn:lsid:zoobank.org:act:F5215674-D97A-4952-AEF5-53CE4AFD983C

Type material. Holotype: 1  ♂, BOLIVIA, Nor Yungas Province, Coroico, 16°12’26”S, 67°43’54”W, elevation ca. 1650 m, mining larva on Liabum sp., Asteraceae, 7–12.vi.2018, ex pupa vi.–vii.2018, field card no. 5239, leg. A. Diškus, J.  R.  Stonis, genitalia slide no.  AD1085 (ZIN). Paratypes (16  ♂, 18  ♀): 6  ♂, 7  ♀, BOLIVIA, same label data as holotype, genitalia slide no. AD1084♀ (ZIN). Paratypes: 1 ♂, 1 ♀, same label data as holotype (NRC); 4 ♂, 6 ♀, PERU, Urubamba Province, near Machu Picchu, 13°9’35”S, 72°31’58”W, elevation ca. 2010 m, mining larvae on Liabum  sp., Asteraceae, 19.x.2008, ex pupa xi.2008, field card no. 4939, leg. A. Diškus, genitalia slides nos. AD842♂, AD1080♂, AD886♀, AD1081♀ (ZIN); 5  ♂, 3  ♀, PERU, Urubamba Province, near Machu Picchu, 13°9’44”S, 72°32’29”W, elevation ca. 2190 m, mining larvae on Smallanthus sp., 19.x.2008, ex pupa xi.2008, field card no. 4943, leg. A. Diškus, genitalia slides nos AD844♂, AD1078♂, AD1079♀ (ZIN); 1 ♀ same label data (NRC).

Diagnosis. From the most similar Neotischeria longa sp. nov. and N. ochripennata sp. nov. (described below) it differs in the smaller size of the adults and male genitalia. Additionally, from N. longa sp. nov. it differs in the white leaf mines (dark brown in N. longa); from N. ochripennata it differs in the blackish brown darkenings on male forewing (forewing ochre-yellow in N. ochripennata) and the slender phallus (basally widened in N. ochripennata).

Male (Figs 34–36). Forewing length 3.3–3.9 mm; wingspan 7.6–8.4 mm (n = 14).

Head. Frons, palpi, and pectens ochreous cream; frontal tuft and collar comprised of brownish grey, cream-tipped lamellar scales; sometimes collar grey-black; antenna significantly longer than one half the length of forewing; flagellum dark grey-brown, with the first antennal segment ochreous yellow; at certain angle of view, flagellum ochre cream basally; sensilla rather long, distinctive.

Thorax. Tegula densely covered with grey-brown, cream-tipped scales or entirely dark grey-black. Thorax glossy, pale brownish grey to dark grey. Forewing dark yellow-ochre, densely speckled with grey-brown or blackish grey brown scales along the costal margin and apically; fringe grey to dark grey, without a fringe line; forewing underside dark brown-grey to black-grey, with small scaleless spots and grey-black special scales along costa basally. No androconia. Hindwing grey to dark grey on upper side and underside, without androconia; fringe grey. Legs pale yellow-ochre, dark brown-grey to black-grey on the upper side.

Abdomen. Glossy, black-grey or grey, sometimes with some purple iridescence on upper side; paler, glossy grey, usually with some ochre scales on underside; anal tufts dorsal, long, dark grey but not always distinctive; genital segments dark grey to grey. Genitalia (Figs 50, 55–59) with capsule 420–430 µm long (or 435– 500  µm long if measured from vinculum to valval tips), 225–250 µm wide. Uncus (Figs 50, 58) comprised of two long latero-ventral lobes and two short rounded dorsal lobes (Fig. 56). Valva (Figs 55, 58) 300–350 µm long (excluding the  basal process), undivided, slender, basally fussed with anellus (Figs  50, 55). Anellus (Fig. 55) strongly thickened laterally, with two long, membranous lobes caudally. Vinculum triangular (Figs 50, 55, 57). Phallus about 375 µm long, slender, apically divided, without lobes or spines (Figs 57, 59).

Female (Figs 38–42). Forewing length 2.9–4.0 mm; wingspan 6.5–8.7 mm (n = 13).

Head. Similar to male but tends to be darker.

Thorax. Usually darker than in male. Forewing with large, irregular, ochre spots.

Abdomen. Genitalia (Figs  53, 54, 81–86) about 1465 µm long. Ovipositor lobes (Fig. 85) relatively large, rounded, densely clothed with peg-like setae; area between ovipositor lobes relatively wide, with tiny papillae usually collected in two rows (Fig.  53). Second pair of lobes, lateral and anterior to the  ovipositor lobes, twice smaller than ovipositor lobes and bearing very long slender lamellar and piliform setae. Anterior and posterior apophyses almost equal in length; prela comprised of three pairs of rod-like projections (Fig. 86). Corpus bursae (Fig.  81) relatively short, basally oval-shaped, without pectinations or signum. Accessory sac absent or indistinctive. Ductus spermathecae very slender, with about 8–10 very large coils (Figs 82, 83).

Bionomics (Figs 8–17). Host plants (Figs 8, 9, 13, 14) are Asteraceae plants: Liabum  spp. (Figs 8, 9, 13, 14), including L. vargasii H. Rob. or L.  solidagineum (Kunth) Less. and Smallanthus sonchifolius (Poepp.) H. Rob. (Fig. 15). Larvae mine in leaves in June and October and produce blotch-like leaf mines (Figs  10, 11, 15–17). Pupation in a circular distinctive nidus inside the leaf mine (Figs 10, 15, 17); colour of the nidus varies from transparent white to pale brown. Adults fly in late June–July and November. Otherwise, the biology is unknown.

Distribution. The  species is known from the tropical or subtropical habitats in the eastern part of the Peruvian and Bolivian Andes, at an elevation of about 1600–2200 m.

Etymology. The  species name is derived from the Latin conexa (combined), in reference to the combined rearing samples from different host plants.

2. Neotischeria ochripennata Diškus & Remeikis, sp. nov.

urn:lsid:zoobank.org:act:3929E91F-D190-4613-A9FC-9ECDBFB832DA

Type material. Holotype: 1  ♂, ECUA-DOR: Chimborazo Province, 15 km NE Bucay (Cumanda), 2°11’5”S, 79°4’14”W, elevation ca. 810 m, mining larva on Liabum sp., Asteraceae, 18.ii.2007, field card no. 4871, leg. A. Diškus, J. R. Stonis, genitalia slide no. RA1090♂ (ZIN). Paratypes (2 ♂): 1 ♂, same label data as holo-type, genitalia slide no. AD1086♂ (ZIN); 1 ♂, ECUADOR: Pichincha Province, 31  km NW Alóag, 0°26’37”S, 78°42’57”W, elevation ca. 2220  m, from feeding larva, 25.ii.2007, field card no. 4898, leg. A. Diškus, J. R. Stonis, genitalia slide no.  AD1089♂ (ZIN) (also see Remarks).

Diagnosis. From the  most similar Neotischeria conexa sp. nov. and N. longa sp. nov. (described in the  current paper) it differs in the  bright ochre-yellow male forewing and fringe (fringe grey and forewing densely speckled with dark scales in N. conexa and N. longa). In the male genitalia, N. ochripennata sp. nov. differs in the distinctive, clearly visible basal extensions of the valva (Figs 49, 62, 67). Additionally, N.  ochripennata differs from N.  longa in the white leaf mines (dark brown in N. longa).

Male (Fig. 37). Forewing length 3.8–4.2 mm; wingspan 8.5–9.5 mm (n = 2).

Head. Frons, palpi, and pectens ochre cream to pale ochre-yellow; frontal tuft and collar glossy, comprised of golden cream or pale ochre-yellow lamellar scales; antenna slightly longer than one half the  length of forewing; flagellum pale yellowish grey, basally ochre-yellow; sensilla whitish cream, rather distinctive but very fine.

Thorax. Tegula covered with ochre-brown, cream-tipped scales, distally cream. Thorax ochre cream, speckled with some ochre-brown scales. Forewing bright ochre-yellow, sparsely speckled with some pale brown scales in apical third; fringe bright ochre-yellow to pale ochre-yellow, with a fringe line of brown-black scales; forewing underside dark ochre-brown, with small scaleless spots and brown-black special scales along costa basally. No androconia. Hindwing grey to pale grey-brown on upper side, pale grey on underside, without androconia; fringe pale grey, ochre glossy. Legs ochre-yellow on underside, dark grey-brown on upper side, with or without some purple iridescence.

Abdomen. Golden ochreous yellow, densely speckled with dark ochre-brown scales on upper side and underside; anal tufts lateral and dorsal (the latter longer), pale ochreous yellow; genital segments golden glossy, ochreous cream. Genitalia (Figs 49, 60–67) with capsule about 490 µm long (or 570 µm long if measured from vinculum to valval tips), 245–250  µm wide. Uncus comprised of two long latero-ventral lobes (Fig. 60) and two short, rounded dorsal lobes (Fig. 61). Valva (Figs 62, 66) about 390 µm long (excluding the basal process), undivided, slender, basally fussed with anellus (Fig. 66) and with a distinctive basal extension (Figs  62, 67). Anellus (Fig.  63) long, strongly thickened laterally, with two long, membranous lobes caudally. Vinculum triangular distally (Figs 66, 67). Phallus about 450 µm long, basally widened (Fig.  67), apically divided, without spines (Figs 62–67).

Female. Unknown (see Remarks).

Bionomics (Figs  18–27). Host plant is Liabum floribundum Less. (or L.  stipulatum Rusby), Asteraceae (Figs 26, 27). Larvae mine in leaves in February and produce blotch-like leaf mines (Figs 18–25). Pupation in a circular nidus inside the leaf mine (Fig. 20); the nidus is transparent, rather indistinctive. Adults fly in March. Otherwise, the biology is unknown.

Distribution. The  species is known from tropical and subtropical western part of the equatorial Andes (Ecuador), at an elevation of about 810–2220 m.

Etymology. The  species name is derived from the  Ancient Greek ochra (ochre colour) combined with the Latin pennatus (winged), in reference to the bright, ochre-yellow colour of the forewing.

Remarks. One female specimen with a label as the holotype (genitalia slide no. RA1089, NRC) was studied and photographed, but excluded from the type series. Because the dark scaling and small size of the specimen and our doubts about its belonging to N. ochripennata, this specimen was left unpublished here.

3. Neotischeria longa Diškus & Stonis, sp. nov.

urn:lsid:zoobank.org:act:9FB2AC43-4197-41DD-9174-B92BD5D1385B

Type material. Holotype: 1 ♂, ECUADOR, Chimborazo Province, 30  km NE Pallatanga, 1°52’41”S, 78°54’11”W, elevation ca. 3020  m, from feeding larvae on Liabum sp., 21.ii.2007, field card no. 4877, leg. A. Diškus, J. R. Stonis, genitalia slide no. AD1082♂ (ZIN). Paratypes (4 ♂, 4 ♀): 3 ♂, 3 ♀ same label data as holotype, genitalia slides nos AD843♂, AD1083♀ (ZIN). Paratypes: 1 ♂, 1 ♀, same label data as holotype (NRC).

Diagnosis. From the most similar Neotischeria conexa sp.  nov. (described above) it differs in the  larger size of the  adults and male genitalia. Additionally, from the  central Andean N. conexa sp. nov., the equatorial Andean N. longa differs in the dark brown leaf mines (white in N.  conexa); from the  similar equatorial Andean N.  ochripennata it differs in the  blackish brown darkenings on male forewing (forewing ochre-yellow in N.  ochripennata) and the slender phallus (basally widened in N. ochripennata).

Male (Fig. 43). Forewing length 3.7–5.0 mm; wingspan 8.0–10.8 mm (n = 5).

Head. Frons, palpi, and pecten yellowish cream to pale ochreous yellow or pale ochre; frontal tuft pale brown basally, pale ochreous yellow distally; collar large, comprised of pale brown and brown, ochretipped scales; sometimes collar comprised of pale grey, cream tipped scales; antenna slightly longer than one half the length of forewing; flagellum pale grey-brown, basally ochre cream; sensilla very fine but distinctive.

Thorax. Tegula grey-brown, distally pale ochre-brown. Thorax brownish cream to yellow cream. Forewing yellow-ochre, densely speckled with blackish brown scales in apical half; fringe mostly grey but apically pale yellowish ochre, with a fringe line of blackish brown scales; forewing underside grey-brown to dark grey-brown, with small scaleless spots and brown-black special scales along costa basally. No androconia. Hindwing dark grey on upper side, grey-brown on underside, without androconia; fringe pale grey, ochre glossy. Legs ochre cream, dark brown-grey on upper side.

Abdomen. Ochre-brown on upper side; paler, yellow-ochre, speckled with brown scales on underside; anal tufts dorsal, pale yellow-ochre; genital segments pale yellow-ochre. Genitalia (Figs 48, 68–76) with capsule about 610 µm long (or 690 µm long if measured from vinculum to valval tips), 280–290  µm wide. Uncus comprised of two long lateroventral lobes (Fig. 68) and two short, rounded dorsal lobes (Fig.  69). Valva (Fig.  71) about 460  µm long (excluding the basal process), undivided, slender, basally fussed with anellus (Figs  68, 71). Anellus (Figs  72, 73) strongly thickened laterally, with two long, membranous lobes caudally. Vinculum triangular but rounded distally (Figs  68, 76). Phallus (Fig.  75) about 485  µm long, slender, apically divided (Fig. 74), without lobes or spines.

Female (Fig.  44). Forewing length 4.0–5.4 mm; wingspan 8.8–11.3 mm (n = 4).

Thorax. Similar to male but forewing with bright yellow-ochre spots, thorax brown.

Abdomen. Bright yellow-ochre on underside. Genitalia (Figs 51, 52, 87–90) about 2110 µm long. Ovipositor lobes (Figs 51, 88) very large, rounded, densely clothed with peglike setae; area between ovipositor lobes moderately wide, with tiny papillae usually not collected in two rows (Fig. 51). Second pair of lobes, lateral and anterior to the ovipositor lobes, very small, indistinctive. Anterior apophyses slightly shorter than posterior ones; prela comprised of three pairs of rod-like projections (Figs 87, 89, 90). Corpus bursae (Fig. 87) very long and slender, basally wider, without pectinations or signum. Accessory sac absent or indistinctive. Ductus spermathecae very slender, with about 10 very large coils.

Bionomics (Figs  28–33). Host plant is Liabum sp., possibly L. dillonii D.  G. Gutiérrez & Katinas or L.  igniarium (Humb. & Bonpl.) Less., Asteraceae (Figs 28, 29). Larvae mine in leaves in February and produce dark brown, irregular blotch-like leaf mines (Figs 29–33). Pupation in a circular, indistinctive nidus inside the leaf mine. Adults fly in March. Otherwise, the biology is unknown.

Distribution. The  species is known from the  subtropical western part of the  equatorial Andes (Ecuador), at an elevation of about 3020 m.

Etymology. The species name is derived from the Latin longus (long), in reference to the very large size of the adults and their genitalia.

4. Neotischeria guarani (Diškus & Stonis, 2020) (comb. nov.)

Paratischeria guarani Diškus & Stonis, in Stonis et al., 2020b: 127–129, 142.

Host plant. Elephantopus mollis Kunth (Asteraceae).

Distribution. This species is known from the single locality in Paraguay, Departamento de Itapúa, Hohenau.

Remarks. Neotischeria guarani was illustrated by Stonis et al., 2020b: Figs 44–45 (adult), Figs 92–94 (male genitalia), Figs 95–97 (female genitalia).

5. Neotischeria mesoamericana (Diškus & Stonis, 2020) (comb. nov.)

Paratischeria mesoamericana Diškus & Stonis, in Stonis et al., 2020b: 129, 130–132, 134, 143, 144.

Host plant. Montanoa hibiscifolia Benth. (Asteraceae).

Distribution. Guatemala.

Remarks. Neotischeria mesoamericana was illustrated by Stonis et al., 2020b: fig. 46 (adult), Figs 98–104 (male genitalia), Figs 105–110 (female genitalia).

6. Neotischeria belizensis (Remeikis & Stonis, 2020) (comb. nov.)

Paratischeria belizensis Remeikis & Stonis, in Stonis et al., 2020a: 64–66.

Host plant. Unknown.

Distribution. This species is known from the  single locality in Belize, Chiquibul Forest Reserve, Las Cuevas.

Remarks. Neotischeria belizensis was illustrated by Stonis et al., 2020a: Figs 24–25 (adult), Figs 124–127 (male genitalia).

7. Neotischeria robinsoni (Diškus & Stonis, 2020) (comb. nov.)

Paratischeria robinsoni Diškus & Stonis, in Stonis et al., 2020a: 55–59.

Host plant. Otopappus verbesinoides Benth. (Asteraceae).

Distribution. This species was described from Belize but recently five specimens (3 ♂, 2  ♀)have also been discovered in Guatemala (ZIN).

Remarks. Neotischeria robinsoni was illustrated by Stonis et al., 2020a: Figs 18–20 (adult), Figs 90–106 (male and female genitalia).

8. Neotischeria tubifex (Diškus & Stonis, 2020) (comb. nov.)

Paratischeria tubifex Diškus & Stonis, in Stonis et al., 2020a: 59–64.

Host plant. Lasianthaea fruticosa (L.) K. M. Becker (Asteraceae).

Distribution. This species is known from the  single locality in Belize, Chiquibul Forest Reserve, Las Cuevas.

Remarks. Neotischeria tubifex was illustrated by Stonis  et  al., 2020a: Figs  21–23 (adult), Figs  107–117 (male genitalia), Figs  118–123 (female genitalia).

The neotropicana group

9. Neoticheria neotropicana (Diškus & Stonis, 2015) (comb. nov.), a type species of the genus

Astroticheria neotropicana Diškus & Stonis, 2015: 457–465.

Paraticheria neotropicana (Diškus & Stonis), in Stonis et al., 2017: 14; 2020a: 66–68; 2020b: 135, 147.

Host plants. Sida rhombifolia L., S. spinose L., and S. acuta Burm.f. (Malvaceae).

Distribution. Paratischeria neotropicana is a tischeriid species with the broadest distribution range yet documented in the  Neotropics, having been recorded from various localities in Belize, Mexico, Guatemala, Ecuador, Peru and Bolivia (Diškus, Stonis, 2015; Stonis et al., 2020b).

Remarks. Neotischeria neotropicana was illustrated by Diškus, Stonis, 2015: figs: 12–13 (adult), Figs 14–21 (male genitalia), Figs 22–27 (female genitalia).

10. Neotischeria capnota (Meyrick, 1915) (comb. nov.)

Tischeria capnota Meyrick, 1915: 247.

Paratischeria capnota (Meyrick, 1915), Xu et al., 2017: 331.

Host plant is unknown.

Distribution: Peru (Lima).

Remarks. Male genitalia of the lectotype and paralectotype (deposited at the NHMUK) was studied and documented for the first time (see Figs 91–98).

11. Neotischeria poseidonia Diškus & Stonis, sp. nov.

urn:lsid:zoobank.org:act:B128F35A-A762-47A7-8457-5C2357C4ABDF

Type material. Holotype: ♂, GUATEMALA: Antigua Guatemala, San Juan del Obispo, 14°31’7”N, 90°43’50”W, elevation ca.  1680  m, from feeding larva, 25.ii.2012, field card no. 5113, A. Diškus, genitalia slide no. AD1090♂ (ZIN).

Diagnosis. Neotischeria poseidonia sp. nov. belongs to the  N.  neotropicana group. In the male genitalia, N. poseidonia from the most similar N.  neotropicana and all other congeneric species differs in the strongly developed, broadly bifurcated and wide phallus (Fig. 80), caudally well-thickened vinculum (Fig.  78), and the distally membranous anellus (Fig. 79).

Male (Figs  45–47). Forewing length about 3.5 mm; wingspan about 7.7 mm (n = 1).

Head. Frons and palpi yellowish cream; frontal tuft comprised of lamellar scales, glossy yellowish cream; collar comprised of lamellar scales, yellowish cream with some pale grey scales; antenna only slightly longer than one half the  length of forewing; flagellum glossy pale grey with some purple iridescence, basally yellowish cream; sensilla medium long, very fine, rather inconspicuous.

Thorax. Tegula speckled with dark grey scales, distally yellow-ochre; thorax yellow-ochre Forewing golden glossy, cream, densely speckled with pale yellow-ochre scales, with irregular pattern of black scales; fringe pale yellow-ochre, with a fringe line of black scales; forewing underside grey-brown, with brown-black scales along costa margin basally. No androconia. Hindwing and fringe brownish cream to pale grey depending on the angle of view. Legs golden cream, densely speckled with blackish grey scales on upper side.

Abdomen. Grey, with blue and purple iridescence on upper side; ochreous cream, distinctly annulated with dark grey-brown scales on underside; anal tufts dorsal, long, ochreous cream; genital segments ochreous cream. Genitalia (Figs  77–80) with capsule 470  µm long, 200 µm wide. Uncus (Figs 77, 79) comprised of two long and slender lateral lobes. Valva (Fig. 79) about 340 µm long (excluding the basal process), slender, undivided, basally fused with anellus (Fig. 79). Anellus thickened only laterally, distally membranous, comprised of two indistinctive membranous lobes. Vinculum triangular but widely rounded distally (Figs 78, 79). Phallus 360 µm long, very wide, robust, broadly bifurcated distally, without spines (Fig. 80).

Female. Unknown.

Bionomics. Host plant is unknown; it is expected to be a  Malvaceae plant. Adults fly in March. Otherwise, the biology is unknown.

Distribution. The  species is known from the single locality in Guatemala (Antigua Guatemala), at an elevation of about 1700 m.

Etymology. The  species name poseidonia (feminine, arbitrary created) is derived from Poseidon, one of the Twelve Olympians, a god of the sea, earthquakes, and horses in the Ancient Greek mythology, in reference to the colossal, fork-like phallus in the male genitalia of Neotischeria poseidonia sp. nov.

Note. Two other new Neotischeria species from Guatemala are already recognized but still undescribed (Stonis  et  al., in prep.). They do not belong to the neotropicana group.

DISCUSSION

The majority of the  currently known Neotischeria species, along with the majority of Astrotischeria, are trophically associated with Asteraceae, and only some utilize plants from Malvaceae. It is not known why Tischeriidae have been so successfully utilizing Asteraceae in the  Americas (Stonis  et  al., 2020b). However, the estimation of the richness and taxonomic diversity of Asteraceae by Katinas et al. (2007) indicates that Central and South America are characterised by the  globally highest generic diversity. Moreover, according to Barreda et al. (2010, 2012), the earliest fossils confidently assigned to Asteraceae suggest a South American-Antarctican origin of the Asteraceae family.

Recently it was shown that plants from Asteraceae are the most diverse hosts for many species of Stigmella Schrank, Nepticulidae in South America (Stonis et al., 2018b), and important host plants for other lepidopteran leaf miners (e.g. Vargas et al., 2012). It is surprising, but there are no Asteraceae-feeding Tischeriidae outside the Americas.

The new genus appears to be almost endemic to Central and South America (the Neotropical region) with a couple of species discovered further to the  North: Arizona and California (USA). The Liabum host plants of the species described in this paper are characterised by a  specific distribution from Central America to the  central Andes (see Gutiérrez, Katinas, 2015) or, in the case of Smallanthus, to the Atlantic Coast of Brazil (see Vitali et al., 2015).

Two other species of this genus have been dissected, studied, and recognised, but not published yet (Stonis et al., in prep). We expect that this genus may comprise almost the double number of species compared with 13 provided in the checklist. Most likely, more species of Neotischeria will be discovered in future in the  heavily undersampled Brazil and perhaps all other areas.

ACKNOWLEDGEMENTS

We are indebted to our Ecuadorian scientific partners, including those who participated in the  initial project in 1999 by R.  Puplesis and S.  R.  Hill, with Professor Giovanni Onore, a  former professor at the  Pontifical Catholic University of Ecuador, Quito, Ecuador. We thank Prof. Dr Jack Shuster (Universidad del Valle de Guatemala, Guatemala City, Guatemala) and the Consejo Nacional de Areas Protegidas (CONAP) Guatemala, C.A. for the Licencia de Collecta o Aprovechamiento de Vida Silvestre (No. 12900). Jonas R. Stonis thanks Julia Puplesyte-Chambers and the  Environmental Programme at the Andes Office of NGO DAR Peru for permission to provide training courses and fieldwork within the project ‘Rapid assessment of biodiversity plots of critical value in the provinces Chanchamayo and Satipo, Peru, and Bolivia’, in cooperation with the  Baltic-American Biotaxonomy Institute in 2017–2018. We thank Prof. Dr Owen T. Lewis (University of Oxford, United Kingdom) for the  material that he collected in 1997–1998 at Las Cuevas, a research station established jointly by the Forest Department of Belize and the Natural History Museum, London in 1994.

For the  help with the  host plant identifications, we are also grateful to Dr Diego G.  Gutiérrez (Museo Argentino de Ciencias Naturales), the author of the so useful monograph Systematics of Liabum Adanson (Asteraceae, Liabeae).

This research was partially funded by a grant (S-MIP-19-30, ‘DiagnoStics’) from the  Research Council of Lithuania.

References

Jonas Rimantas Stonis, Arūnas Diškus, Andrius Remeikis, Brigita Paulavičiūtė, Liliana Katinas, Nixon Cumbicus Torres

DIFERENCINĖ PARATISCHERIA IR NEOTISCHERIA GEN. NOV. DIAGNOSTIKA IR NAUJOS, ASTERACEAE ŠEIMOS AUGALUS MINUOJANČIOS, TISCHERIIDAE (LEPIDOPTERA) RŪŠYS IŠ CENTRINĖS BEI PIETŲ AMERIKOS