Zootaxa 3737 (1): 001–023 ISSN 1175-5326 (print edition) Article ZOOTAXA www.mapress.com/zootaxa/ Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3737.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:BF23F07F-4EBC-4309-9504-5AEF8ACDCFB4 First discovery of Quercus feeding Nepticulidae (Lepidoptera) in Central America JONAS R. STONIS1,2,5, ARŪNAS DIŠKUS1,2, ANDRIUS REMEIKIS1, 3 & JACK SCHUSTER4 1Department of Biology, Lithuanian University of Educational Sciences, Studentų St.39, Vilnius LT-08106, Lithuania 2Institute of Public Administration, Mykolas Romeris University, Valakupių St.5, Vilnius LT-10101, Lithuania 3Nature Research Centre, Akademikos St. 2, Vilnius LT-08412, Lithuania 4Systematic Entomology Laboratory, Universidad del Valle de Guatemala, Guatemala City, Guatemala. 5Corresponding author. E-mail: [email protected] Abstract Despite the high taxonomic diversity of oaks in Mexico and Central America, no Quercus feeding Nepticulidae have ever been recorded from the region. Here, we present seven species whose larvae are leaf-miners of Quercus (section Lobatae) in Guatemala. Except Stigmella nigriverticella (Chambers 1875), which was previously known from the United States, all other discovered species are new. We describe and name five new species (Stigmella jaguari Remeikis & Stonis, sp. nov., S. lauta Diškus & Stonis, sp. nov., S. sublauta Remeikis & Stonis, sp. nov., S. aurifasciata Diškus & Stonis, sp. nov. and S. guatemalensis Diškus & Stonis, sp. nov.); the remaining new species is described but left unnamed because of lack of adults (i. e. moths and genitalia are described from developed pupae). All seven treated species are illustrated with pho- tographs of the leaf-mines, adults, and genitalia. Key words: Central America, Guatemala, leaf-mines, Nepticulidae, new species, Quercus, Stigmella Introduction Oaks (Quercus), with about 600 species, are native to the northern hemisphere, extending from temperate to tropical latitudes in South East Asia and the Americas, with greatest taxonomic diversity and most important centres of endemism in South East Asia and North America (particularly southeastern United States, southern Mexico and the montainous part of Central America) (Menickiy 1984, Nixon 2006, Torres-Miranda et al. 2011). In contrast to Europe and East Asia, where leaf-mining or bark-mining Nepticulidae have extensively been studied (e. g. Johansson et al. 1990, Puplesis 1994, Puplesis & Diškus 2003, van Nieukerken & Liu 2000, Nieukerken & Johansson 2003, Stonis & Rocienė 2013, Rocienė & Stonis 2013), Central America has been neglected, with no Quercus feeding nepticulid species known. This paper is the first to report the Central American Nepticulidae species associated with Quercus as a host-plant. Material and methods Adults of all species were collected by the Lithuanian-Guatemalan scientific expedition’ 2012 in montane forests of Guatemala (Quetzaltenango, or more commonly known as Xela, 14°47'27"N, 91°32'09"W; Santa Cruz del Quiché, 15°01'22''N, 91°10'17''W; Panajachel, 14°45'06''N, 91°09'43''W) at elevations ca 1655–2500 m (Figs 2–6) by rearing imagos from mining larvae using the standard method (Puplesis 1994 and Diškus & Stonis 2012). Collecting methods, techniques for genitalia preparation, and protocols for description are outlined in Puplesis & Robinson (2000), Puplesis & Diškus (2003) and Diškus & Stonis (2012). Permanent slides were photographed and studied using a Leica DM2500 microscope and Leica DFC420 digital camera. Accepted by J.F. Landry: 29 Oct. 2013; published: 18 Nov. 2013 1 The descriptive terminology of morphological structures follows van Nieukerken (1985, 1986), Johansson et al. (1990), Puplesis & Robinson (2000) and van Nieukerken & Johansson (2003). The classification of Nepticulidae follows Puplesis (1994), Puplesis & Robinson (2000), Puplesis et al. (2002) and Puplesis & Diškus (2003). Abbreviation for specimen depository: ZMUC Zoological Museum, University of Copenhagen, Denmark. Descriptive taxonomy All Quercus feeding species discovered in Central America (Guatemala) belong to the genus Stigmella Schrank and fall into two species groups: the saginella-group and the quercipulchella-group (diagnostics of the species groups associated with Quercus as a host-plant are extensively discussed in Stonis et al. 2013). Only one of the species discovered, S. nigriverticella (Chambers) described from the United States, was previously known; it is redescribed below from specimens that we collected in Guatemala. All remaining species from Guatemala are new and they are described below. The saginella group Stigmella nigriverticella (Chambers, 1875) (Figs 1, 7–12, 30, 32–34, 46–54) Material examined. 6♂, 14♀, Guatemala, Quetzaltenango, 14°47'27"N, 91°32'09"W, mining larvae on Quercus spp., 2500 m, 08.ii.2012, field card no. 5120, LT-GT Scientific Expedition, genitalia slides nos RA483, RA484, RA485, RA486, RA487, RA488, RA490 (ZMUC); 2♂, Panajachel, 14°45'06''N, 91°09'43''W, elevation ca. 1657 m, montane forest, larva on Quercus spp., 22.ii.2012, ex pupa iii.2012, LT-GT Scientific Expedition, genitalia slides nos AD516, AD514 (from pupa) (ZMUC). Diagnosis. The male genitalia differ from Stigmella castaneaefoliella (Chambers 1875) and S. flavipedella (Braun 1914) by the absence of cornuti. From all other species of the saginella group, it differs by the combination of elaborated gnathos, narrow uncus, relatively slender valva and X-shaped juxta. Male (Figs 32–34). Forewing length 2.2–3.0 mm; wingspan 4.8–6.4 mm. Head: palpi cream to yellowish cream; frontal tuft always black on vertex, sometimes yellowish cream in frontal part; collar and scape yellowish cream, occasionally greyish cream; antenna with 27–30 segments, slightly longer than half of forewing; flagellum grey-brown or blackish on upper side, grey-cream to grey on underside. Thorax yellowish cream; tegulae black in front. Forewings yellow-cream with black markings (the pattern may considerably vary; see figs 32, 33); cilia grey to fuscous (on tornus) to cream (on apex); underside of forewing usually blackish, sometime grey-cream to dark grey, but always with elongated yellow-cream spot on forewing apex (Fig. 34). Hindwings and cilia of hindwings vary from usually grey to dark grey. Legs dark grey or black on upper side, yellowish cream on underside. Abdomen black to dark grey on upper side, yellow-cream on underside; anal tufts cream to pale grey. Female. Antenna with ca. 26–28 segments. Abdomen vary from yellow-cream to blackish grey on upper side, yellow-cream on underside; anal tufts cream to yellowish cream. Otherwise as male. Male genitalia (Figs 46–50). Capsule longer (250–270 μm) than wide (200–220 μm). Vinculum without lateral lobes, ventral plate short (35–40 μm). Uncus with short thickened lateral lobes. Gnathos with two caudal, 35–40 μm long processes (Fig. 49). Valva (Figs 47, 48) 200–205 μm long, with numerous long chaetae; apical process weakly individualized; transtilla with very short sublateral processes. Juxta X-shaped (Fig. 48). Aedeagus (Fig. 50) 170 μm long; vesica without cornuti. Female genitalia (Figs51–54). Total length 1055 μm. Apophyses anteriores (about 110 μm) much shorter than apophyses posteriores (about 145μm) (Fig. 52). Vestibulum relatively narrow, without sclerites. Accessory sac very small. Corpus bursae oval, 590–600 μm, covered with numerous pectinations and with long (360–375μm), strongly chitinized signa (Figs 53, 54). 2 · Zootaxa 3737 (1) © 2013 Magnolia Press STONIS ET AL. FIGURES 1–6. Major collecting localities and distribution of the species discovered in Guatemala. 1, distribution of Stigmella nigriverticella (Chambers); 2, distribution of the new Stigmella species; 3, Quetzaltenango, 14°47'27"N, 91°32'09"W, montane mixed forests at elevation 2500 m; 4, same, moist broadleaf tropical forest at elevation 2000 m; 5, Santa Cruz del Quiché, 15°01'22''N, 91°10'17''W, elevation ca 2020 m; 6, Panajachel, 14°45'06''N, 91°09'43''W, elevation ca. 1660 m (Maps – courtesy of T.Patterson, USA). CENTRAL AMERICAN NEPTICULIDAE Zootaxa 3737 (1) © 2013 Magnolia Press · 3 FIGURES 7–12. Bionomics of Stigmella nigriverticella (Chambers). 7–9, leaf-mines; 10, major host-plant Quercus crassifolia Humb. & Bonpl.; 11, other host-plant Q. crispipilis Trel; 12, habitat, montane mixed forest at elevation 2500 m. Bionomics. Mines in leaves (Figs 7–9). Host-plants: Quercus spp., incl. Q. crassifolia Humb. & Bonpl. and Q. crispipilis Trel. (section Lobatae) in Guatemala (Figs 10, 11). Egg on upper side of the leaf. Larvae mine in February. Sinuous gallery of mine filled with blackish frass (Figs7–9). Larva pale green, with brownish intestine. Larval exit slit on upper side of the leaf. Cocoon (Fig. 30) whitish; length 2.4 mm, maximal width 1 mm. Adults emerged in March. 4 · Zootaxa 3737 (1) © 2013 Magnolia Press STONIS ET AL. Distribution (Figs 1, 2). USA (from Cincinnati, Ohio and Pennsylvania to Texas) (Newton & Wilkinson 1982); Guatemala in montane mixed forests at elevation 1655–2500 m (Figs 3, 12). Note. The specimens from Guatemala strongly vary in head markings (frontal tuft from fully black to yellowish cream in frontal part) and forewing markings (see figs 32, 33). However, the male and female genitalia fully correspond with those illustrated in Newton & Wilkinson (1982). FIGURES 13–17. Bionomics of Stigmella jaguari Remeikis & Stonis, sp. nov. 13–15, leaf-mines (a—old mine, b — with feeding larva); 16, host-plant Quercus crispipilis Trel; 17, habitat, montane mixed forest at elevation 2500 m. CENTRAL AMERICAN NEPTICULIDAE Zootaxa 3737 (1) © 2013 Magnolia Press · 5 Stigmella jaguari Remeikis & Stonis, sp.nov. (Figs 2, 13–17, 31, 35–37,55–59) Type material. Holotype: ♂, Guatemala, Quetzaltenango, 14°47'27"N, 91°32'09"W, mining larvae on Quercus sp., 2500 m, 08.ii.2012, field card no. 5119, LT-GT Scientific Expedition, genitalia slide no. RA491 (ZMUC). Paratype: 1♂, label data as holotype, genitalia slide no. RA489 (ZMUC). Diagnosis. In male genitalia the new species differs from all other species of the saginella group except Stigmella lauta sp. nov. by the long vinculum and rounded juxta. From related S. lauta sp. nov. it differs by the short cornuti, bulged valva, and tiny sublateral processes of transtilla (S. lauta posseses long cornuti, stright valva, and large sublateral processes of transtilla). Male (Figs 35–37). Forewing length 2.2–2.5 mm; wingspan 4.8–5.4 mm. Head: palpi cream; frontal tuft black; collar and scape large, yellowish cream; antenna with 27–29 segments, slightly shorter than half of forewing; flagellum pale grey to dark grey upper side, grey to cream-grey on underside. Thorax, tegulae and forewings yellow-cream or grey-cream densely but irregularly speckled (irrorated with black scales, no distinct forewing pattern); cilia grey (including forewing apex); underside of forewing black-grey, with cream spot on apex. Hindwings and cilia grey. Legs fuscous on upper side, cream on underside. Abdomen fuscous on upper side, grey- cream on underside, anal plates and tufts cream. Female. Unknown. Male genitalia (Figs 55–59). Capsule longer (250 m) than wide (175 μm). Vinculum without lateral lobes; ventral plate large. Uncus with short, distally chitinized lateral lobes (Figs 55, 57). Gnathos with slender transverse bar and two caudal processes (Fig. 57). Valva (Figs 55, 57) 120 μm long, bulged medially, with pointed apical process; transtilla with short pointed sublateral processes. Juxta rounded caudally (Fig. 55). Aedeagus (Figs 58, 59)175–195 μm long, 60–70 μm broad; vesica with 7–8 large spine like cornuti and some minute cornuti. Bionomics. Mines in leaves (Figs 13–15). Host-plants: Quercus crispipilis Trel. (section Lobatae) (Figs 16, 11). Egg on upper side of the leaf. Larvae mine in February. Sinuous gallery of mine filled with blackish frass (Figs 13, 14). Larva deep green, with dark green or brown-green intestine. Larval exit slit on upper side of the leaf. Cocoon (Fig. 31) whitish; length 2.3 mm, maximal width 1 mm. Adults emerged in March. Distribution (Fig. 2). Known only from Guatemala in montane mixed forests at elevation 2500 m) (Figs 3, 17). Etymology. This species is named after the jaguar, a sacred animal of the Maya people, in reference to the yellow-cream forewing of S. jaguari irregularly speckled with black scales. Stigmella lauta Diškus & Stonis, sp.nov. (Figs 2, 18–23, 38–40, 60–69) Type material. Holotype: ♂, Guatemala, Santa Cruz del Quiché (Utatlán Ruins), 15°01'22''N, 91°10'17''W, elevation ca. 2023 m, montane forest, larva on Quercus sp., 20.ii.2012, ex pupa iii.2012, field card no. 5103, LT-GT Scientific Expedition, genitalia slide no AD505 (ZMUC). Paratypes: 2♂, 2♀, same label data as holotype, genitalia slide nos AD506♂, AD512♂, AD507♀, AD518♀. Diagnosis. In male genitalia the new species differs from all other species of the saginella group by the very long vinculum, rounded sublateral processes of transtilla, and very long spine-like cornuti. Also see Diagnosis of related S. sublauta sp. nov. Male (Figs 38–40). Forewing length 2.2–2.3 mm; wingspan 4.8–5.0 mm. Head: palpi cream; frontal tuft black on vertex, yellowish cream in smaller frontal part; collar and scape large, yellowish cream; antenna with 27–29 segments, slightly shorter than half of forewing; flagellum dark grey on upper side, grey to cream-grey on underside. Thorax, tegulae and forewings yellow-cream or grey-cream densely irrorated with brown-black or black scales (no distinct forewing pattern); sometimes dark scales prevail and forewings look dark brown with a few scattered pale scales; cilia grey to blackish grey (including forewing apex); underside of forewing black-grey, with distinct or indistinct elongated cream spot on apex. Hindwings and cilia grey. Legs fuscous on upper side, yellowish cream on underside. Abdomen fuscous on upper side, cream on underside. Female. Similar to male. 6 · Zootaxa 3737 (1) © 2013 Magnolia Press STONIS ET AL. FIGURES 18–23. Bionomics of Stigmella lauta Diškus & Stonis, sp. nov. 18–22, leaf-mines; 21, host-plant Quercus sp. (section Lobatae); 23, habitat, montane mixed forest at elevation 2020 m. Male genitalia (Figs 60–64). Capsule longer (335 μm) than wide (185 μm). Vinculum without lateral lobes; ventral plate very large. Uncus with short triangular, distally thickened lateral lobes (Fig.62). Gnathos with slender transverse bar and two caudal processes (Figs 60, 62). Valva (Fig.61) 145 μm long, slender, pointed apically; transtilla with broad rounded sublateral processes. Juxta rounded caudally (Figs 60, 61). Aedeagus (Figs 63,64) 214–235 μm long, 55–60 μm broad; vesica with 6 very long spine-like cornuti. CENTRAL AMERICAN NEPTICULIDAE Zootaxa 3737 (1) © 2013 Magnolia Press · 7 Female genitalia (Figs 65–69). Total length 905 μm. Apophyses anteriores slightly shorter than apophyses posteriores (Fig. 67). Vestibulum relatively narrow, without sclerites. Accessory sac very small. Corpus bursae oval, 410 μm, covered with numerous pectinations and with short (55–110 μm) chitinized signa (Figs 68, 69). Bionomics. Mines in leaves (Figs 18–23). Host-plants: Quercus sp. (section Lobatae) (Figs 21). Egg on upper side of the leaf. Larvae mine in February. Sinuous or contorted gallery of mine filled with coiled dark brown or black frass (Figs 18–22). Larva deep green, with brownish green intestine (Fig. 18). Larval exit slit on upper side of the leaf. Cocoon beige-brown. Adults emerged in March. Distribution (Fig. 2). Known only from Guatemala in mixed montane forests at elevation about 2000 m (Fig.23). Etymology. The species name is derived from Latin lautus (elegant; gaudy) in reference to the very large ventral plate of the vinculum, and transtilla with unusually broad, rounded sublateral processes. Stigmella sublauta Remeikis & Stonis, sp. nov. (Figs 2, 3, 24–26, 41, 70–75) Material examined. Holotype: ♀, Guatemala, Quetzaltenango, 14°47'27"N, 91°32'09"W, mining larvae on Quercus crassifolia, 2500 m, 08.ii.2012, field card no. 5118, LT-GT Scientific Expedition, genitalia slide no. RA481 (ZMUC). Paratypes: 2♀, the same label as holotype (ZMUC). Diagnosis. It differs from all other species of the saginella group by combination of speckled forewings, long apophyses anteriores and asymmetrical signa. From most similar S. lauta sp. nov. it also differs by the paler forewing, much longer apophyses anteriores and signa: in S. sublauta signa are 175 and 280 μm long (in S. lauta 55–110 μm). Male. Unknown. Female (Fig. 41). Forewing length 2.3–2.4 mm; wingspan 5.0–5.2 mm. Head: palpi cream; frontal tuft black; collar and scape cream; antenna with 19–20 segments, slightly shorter than half of forewing; flagellum dark grey to fuscous on upper side, grey to cream-grey on underside. Thorax, tegulae and forewings yellow-cream or grey- cream densely irrorated with dark brown or black-brown scales (with no distinct forewing pattern); cilia pale grey; underside of forewing very pale brownish grey, except dorsal and tornal edges and apex where it remains cream. Hindwings and cilia pale grey. Legs fuscous to grey on upper side, yellowish cream on underside. Abdomen pale grey on upper side, yellowish cream on underside. Female genitalia (Figs 70–75). Total length 1050 μm. Anterior and posterior apophyses equally long, ca. 150 μm (Fig. 70). Vestibulum narrow, heavily wrinkled, without sclerites. Accessory sac very small. Corpus bursae elongated, 660 μm, covered with numerous pectinations and with distinctly asymmetrical, 175 and 280 μm, chitinized signa (Figs 73–75). Bionomics. Mines in leaves (Figs 24, 25). Host-plant: Quercus crassifolia Humb. & Bonpl. (section Lobatae) (Fig. 26). Egg on upper side of the leaf. Larvae mine in February. Sinuous gallery of mine filled with blackish frass (Figs 24, 25). Larva yellowish, with brownish intestine. Larval exit slit on upper side of the leaf. Cocoon whitish. Adults emerged in March. Distribution (Fig. 2). Known from Guatemala (mixed montane forests at elevation 2500 m (Fig. 3). Etymology. This species is named after related Stigmella lauta Diškus & Stonis. Stigmella aurifasciata Diškus & Stonis, sp. nov. (Figs 2, 42, 76–80) Material examined. Holotype: ♀, Guatemala, Panajachel, 14°45'06''N, 91°09'43''W, elevation ca. 1657 m, montane forest, larva on Quercus sp., 22.ii.2012, ex pupa iii.2012, LT-GT Scientific Expedition, genitalia slide no AD499 (ZMUC). Diagnosis. It differs from all other species of the saginella group by combination of very broad golden cream median and two black basal and apical fasciae of forewing, very long apophyses posteriores, and asymmetrical signa. 8 · Zootaxa 3737 (1) © 2013 Magnolia Press STONIS ET AL. FIGURES 24–29. Bionomics of Stigmella species. 24, 25, S. sublauta Remeikis & Stonis, sp. nov., leaf-mines; 26, same, host- plant Quercus crassifolia Humb. & Bonpl. (section Lobatae); 27, S. aurifasciata Diškus & Stonis, sp. nov., leaf-mine; 28, same, host-plant Quercus sp. (section Lobatae); 29, same, type locality, 1660 m (Panajachel). Male. Unknown. Female (Fig. 42). Forewing length 2.3 mm; wingspan 5.0 mm. Head: palpi; frontal tuft brownish orange; collar and scape yellow-cream; antenna with about 26 segments, slightly shorter than half of forewing; flagellum fuscous on upper side, cream on underside. Thorax golden cream (or yellow-cream); tegulae slightly darker, yellow-cream. Forewings with very broad golden cream (or yellow-cream) median fascia and two broad black CENTRAL AMERICAN NEPTICULIDAE Zootaxa 3737 (1) © 2013 Magnolia Press · 9 fasciae: basal and apical; tiny basal area before basal fascia remains yellow-cream; fuscous scales with some purple iridescence; cilia yellowish cream on apex, greyish on tornus; underside of most of forewing fuscous, apex yellowish cream. Hindwings pale grey on upper side; grey to dark grey on underside; cilia of hindwings grey. Legs dark grey to fuscous on upper side, yellow-cream on underside. Abdomen black on upper side, yellow-cream on underside. Female genitalia (Figs 76–80). Total length 845 μm. Anterior apophyses shorter than posterior apophyses (Figs 77, 78). Vestibulum narrow, without sclerites. Accessory sac very small. Corpus bursae elongated, 460 μm, covered with numerous pectinations (Fig. 79) and with asymmetrical, 135 and 230 μm, chitinized signa (Figs 76, 80). Bionomics. Mines in leaves. Host-plant: Quercus sp. (section Lobatae). Larvae mine in February. Adults emerged in March. Distribution (Fig. 2). Known only from Guatemala in mixed montane forests at elevation about 1660 m. Etymology. The species name is derived from Latin aureus (golden) and fasciata (with fascia) in reference to the forewing with a broad golden cream fascia. The quercipulchella group Stigmella guatemalensis Diškus & Stonis, sp. nov. (Figs 2, 43–45, 81–84) Type material. Holotype: ♂, Guatemala, Santa Cruz del Quiché (Utatlán Ruins), 15°01'22''N, 91°10'17''W, elev. ca. 2023 m, montane forest, larva on Quercus sp., 20.ii.2012, ex pupa iii.2012, LT-GT Scientific Expedition, genitalia slide no. AD513 (ZMUC). Diagnosis. The new species differs from other species of the quercipulchella group by the large apical process of valva, large triangular process of inner lobe of valva, very long and narrow anterior processes of gnathos, and lack of sublateral processes of transtilla. Also see Diagnosis of S. species 515. Male (Figs 43–45). Forewing length 1.8 mm; wingspan 4.0 mm. Head: palpi cream; frontal tuft dark orange; collar and scape cream; antenna with 29 segments, slightly longer than half of forewing; flagellum grey on upper side, pale grey on underside. Thorax, tegulae and forewing fuscous with distinct purple and blue iridescence; fascia of forewing postmedian, broadening at tornus, cream with distinct golden lustre; cilia fuscous; underside of forewing fully black. Hindwings fuscous grey, covered with black androconia (most overlapping over cilia) (Fig. 44); cilia of hindwings blackish grey. Legs dark grey to fuscous on upper side, grey-cream on underside. Abdomen shiny, fuscous, with purple and green iridescence on upper side, silvery grey with some purple and greenish iridescence on under side; genital plates fuscous, with some purple iridescence; anal tufts fuscous, very short, indistinct. Female. Unknown. Male genitalia (Figs 81–84). Capsule longer (255 m) than wide (190 μm). Vinculum with triangular lateral lobes. Uncus distinctly bilobed, laterally thickened (Fig. 82). Gnathos with short caudal processes and very long, narrow anterior processes (Fig. 82). Valva 155 μm long, with very large apical process and large triangular process on inner lobe; transtilla without sublateral processes. Aedeagus (Fig. 84), 240 μm long, 120 μm broad; vesica with numerous spine-like cornuti collected into a long curved band and with an apical group of larger spine-like cornuti; manica present. Bionomics. Mines in leaves. Host-plant: Quercus sp. (section Lobatae). Egg on upper side of the leaf. Larvae mine in February. Sinuous or contorted gallery of mine filled with blackish frass. Larva deep green, with dark green intestine. Larval exit slit on upper side of the leaf. Cocoon ochre-brown. Adults emerged in March. Distribution (Fig. 2). Known only from Guatemala in mixed montane forests at elevation about 2000 m. Etymology. This species is named after Guatemala, a beautiful and megadiverse country from where it was found. 10 · Zootaxa 3737 (1) © 2013 Magnolia Press STONIS ET AL.