Rediscovery of Glenurus Incalis Banks (Neuroptera: Myrmeleontidae), and Notes on the Brazilian Species of Glenurus Hagen

Zootaxa 4858 (1): 135–143 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4858.1.10 http://zoobank.org/urn:lsid:zoobank.org:pub:FFDB3C6B-474F-4581-95B5-07C7AEBE2248

Rediscovery of Glenurus incalis Banks (Neuroptera: Myrmeleontidae), and notes on the Brazilian species of Glenurus Hagen

RENATO JOSE PIRES MACHADO Universidade Federal do Paraná (UFPR), Departamento de Zoologia, Curitiba, Paraná, Brazil. �[email protected]; https://orcid.org/0000-0003-3155-3639

Abstract

The antlion species, Glenurus incalis Banks is the most enigmatic species of this New World genus. It was described almost 100 years ago based on a single female collected in the Peruvian Amazon, which until today remained as the only known specimen. Herein I report four new specimens of Glenurus incalis, which were used to thoroughly redescribe the species and to expand its distribution to Brazil. Notes on the other Brazilian species of Glenurus is also presented, including the first record of G. heteropteryx Gerstaecker to the country.

Key words: antlion, new records, distribution map, lacewings

Introduction

The genus Glenurus Hagen, 1866 is represented by some of the most charismatic antlions in the New World. The adults are relatively large insects (reaching over 90 mm of wingspan) with decorated wings presenting large brown, white and pinkish marks. Different of most antlions the Glenurus larvae do not live burrowed in sand, they are known to habit dry tree holes, under rocks, and even tortoise burrows (Stange 2000; 2004). According to the most recent Myrmeleontidae classification (Machado et al. 2019), Glenurus is the type genus of the New World antlion tribe Glenurini (Myrmeleontidae: Nemoleontinae), and is represented by nine species ranging from USA to Argentina (Stange 2004). However, some previous classifications (Banks 1910; Markl 1954) placed some unrelated Asian species within Glenurus, including some species that are today classified in Dendro- leontinae. Furthermore, there are three Asian species, G. atomatus Yang, 1986 and G. fuscilomus Yang, 1986 from China, and G. posticus Navás, 1913 from Vietnam, that are still equivocally classified in Glenurus (Stange 2004; Oswald 2019; Machado et al. 2019), but they are not considered here, since this classification is clearly inaccurate and these three species should be transferred to a different genus. More recently the Glenurus, as interpreted here, was recovered as monophyletic by Badano et al. (2018), who used three species and morphological data. This result was later reinforced by a different study comprising two species and genomic data, which also recovered the genus monophyletic (Machado et al. 2019). Interest in Glenurus has been increasing in recent years, with a series of papers published on the biology and the distribution records of different species (Stange 2000; Miller & Stange 2006; Ardila-Camacho et al. 2014; Giacom- ino 2015; Petko et al. 2016). Though, none of these papers mentioned, Glenurus incalis Banks (Fig. 1), which was firstly introduced almost 100 years ago. The species was succinctly described (7 lines text) based on a sole female collected in Peru (Banks 1922), and until now the holotype remained as the only known specimen of G. incalis. The study of antlions from different Brazilians institutions revealed some new specimens of Glenurus, including four individuals of G. incalis. In this sense, based on the complete absence of information on this species during the past 100 years, this study aims to thoroughly redescribe G. incalis and also report new distributional records of Glenurus in Brazil, which was previously represented only by G. peculiaris (Walker).

Accepted by A. Letardi: 25 Aug. 2020; published: 29 Sept. 2020 135 Figure 1. Glenurus incalis, dorsal view.

Material and methods

A total of 19 specimens of Glenurus were studied herein. The specimens were obtained from different institutions (see list below) either by visit, loans or high-resolution images. Species were identified by the key provided by Banks (1922) and comparison with images of the type specimens. The images were taken with a stereomicroscope (Leica M205 C) equipped with a digital camera (Leica MC190 HD), using focus stacking software (Leica Appli- cation Suite, version 4.2). Genitalia drawings were vectorized with Adobe Illustrator CS3. Distribution map was prepared in ArcGIS and later edited in Adobe Illustrator CS3. Morphological terminology follows Stange (1994).

List of the insect collections cited

DZUP—Coleção Entomológica Pe. Jesus Santiago Moure, Curitiba, PR, Brazil Embrapa-DF—Embrapa Cerrados, Planaltina, DF, Brazil Esalq—Escola Superior de Agricultura Luiz de Queiroz, Piracicaba, SP, Brazil INPA—Instituto Nacional de Pesquisas da Amazônia, Manaus, AM, Brazil MCZ—Museum of Comparative Zoology, Cambridge, MA, USA MNRJ—Museu Nacional do Rio de Janeiro, Rio de Janeiro, RJ, Brazil MZSP—Museu de Zoologia de São Paulo, São Paulo, SP, Brazil UFBA—Universidade Federal da Bahia, Salvador, BA, Brazil UFLA—Universidade Federal de Lavras, Lavras, MG, Brazil

Results

Among the examined material three species were recognized, G. peculiaris, G. incalis, and G. heteropteryx Ger- staecker, with the last two being recorded from Brazil for the first time. Coincidentally all studied specimens were females, except for two individuals that the tip of the abdomen was lost and the sex was impossible to determine. Details for each of these three species, including an identification key, and the redescription of G. incalis are presented below.

Glenurus peculiaris was the only species of the genus previously known from Brazil. It can be distinguished from the other two species recorded herein to the country by the combination of the presence of a large apical brown mark in the forewing, and the two white spots on the posterior margin of the apical area in the hind wing (Fig. 2c). The species has a large distribution in South America, with records from Argentina, Guyana, Paraguay and Suriname, but in Brazil it was only known from two records from São Paulo state (Stange 2004; 2010; Machado & Martins 2020). These two Brazilian records correspond to the two specimens of the type series of G. brasiliensis Navás, 1920, a junior synonym of G. peculiaris. The holotype of G. peculiaris, despite being collected in Brazil, had no extra location data on the label. Navás (1920) in the description of G. brasiliensis, mentioned that the female specimen was from Jaragua, which was interpreted by Oswald (2019) as the municipality of Jaraguá do Sul in Santa Catarina state. However, Navás (1923) mentioned that this record was erroneous and that the specimen was actually from nearby the municipality of São Sebastião in São Paulo. Herein, I present the first records of G. peculiaris from four different Brazilian states, Bahia, Minas Gerais, Paraná, and Santa Catarina besides some new records from São Paulo state (Fig. 5), all areas dominated by the Atlantic rainforest biome. Examined specimens. Brazil: Bahia: Caeté Açu, Vale do Capão, Lothlorien, 4.xii.2016, manual, Pium, Burg- er, R. (1♀—UFBA); Minas Gerais: Lavras (1? UFLA); Paraná: Foz do Iguaçu, 13.i.1967, Bourlegat (1♀ Esalq); Morretes, Bairro América de Cima, 30.xii.2018, BR. Araujo (1♀ DZUP); Rolandia (4♀ MZSP); Santa Catarina: Joinvile (1? MNRJ—certainly destroyed in the fire of 2.ix.2018); Nova Teutonia, 27º11’S–52º23’W, 300–500m, xii.1976, Fritz Plaumann (1♀ INPA); São Paulo: Caraguatatuba, ii.1963, Ubirajara col. (1♀ MZSP); Registro, 30.xii.1968, R.G. da Silva col. (1♀ MZSP); Ubatuba, Praia Fortaleza 6.iii.1973, Froelich col. (1♀ MZSP).

Glenurus heteropteryx Gerstaecker, 1885

Glenurus heteropteryx can be easily distinguished from any other species on the genus because of its mostly transparent forewing (Fig. 2a), all other eight species present a large apical brown band in the forewing. The species was initially described from Panama and was later mentioned by Stange (2002) as possibly also occurring in Costa Rica. Posteriorly it was recorded from some of the Caribbean islands (Trinidad and Guadeloupe) and also from South America (Ecuador, Colombia, and Venezuela) (Stange 2004; Ardila-Camacho et al. 2014; Giacomino 2015). Herein I report the first record of G. heteropteryx from Brazil, one specimen collected in the northern state of Roraima (Fig. 5). Examined specimens. Brazil: Roraima: Ilha de Maracá, 20–30.iii.1987, Luis S. Aquino (1♀ INPA).

Glenurus incalis Banks, 1922

As mentioned previously, G. incalis is known only by the female holotype from Chanchamayo Peru, but herein I report four new specimens, another one from Peru and other three from the Amazon region in Brazil (Fig. 5). The species is thoroughly redescribed below based on these four specimens and high-resolution images of the holotype. Lengths: forewing: 41.5–44.5 mm; hind wing: 43–47 mm. Head (Fig. 3): Labrum and clypeus mostly dark brown, but lateral margins light brown; set with pale setae. Frons below antennae light brown with medial region dark brown, space between and above the antennae dark brown; set with few short pale setae. Gena light brown. Vertex raised; mostly brown with a medial longitudinal and medial transversal lines dark brown; set with short black setae. Ocular rim setae absent. Antennae clavate; more than three times longer than pronotum length; distance between antennae about the same size of scape width; scape and pedicel brown to dark brown; 49–51 flagellomeres about as long as wide, except by the basal one, about twice longer than wide, and the apical ones, much wider than long; flagellum brown but darkening towards the apex; all segments set with short black setae. Mandible light brown with internal margin and apex black. Maxillary palpi light brown except by the last two distal segments, dark brown. Labial palpi light brown except by the distal segment, dark brown; distal palpomere fusiform, palpimacula oval-shaped, located medially. Thorax (Fig. 3b–c): Pronotum slightly longer than wide; posterior margin wider than anterior; subapical furrow present; mostly dark brown, except by a pale longitudinal stripe medially and the area distal to the furrow, brown

Redescription of Glenurus incalis Zootaxa 4858 (1) © 2020 Magnolia Press · 137 to pale; set with long black setae. Meso and metanotum dark brown, except by a pale longitudinal stripe medially (stripe fading out in the metascutellum and sometimes completely absent, making it entirely dark brown); set with white setae and few black ones at the anterior margin of the mesoprescutum. Pterothoracic pleura entirely dark brown, except by small pale-yellow marks at the base of the wings; set with long white setae, and a few long black setae on the anterior margin of the mesothorax. Wings (Fig. 2b): Elongate and with apex acute. Forewing shorter and wider than hind wing. Anterior Banksian line present in both wings but more evident in the forewing; posterior Banksian line absent in both wings. Sc, RA and Cu veins intercalating dark brown and pale areas, remaining veins entirely dark brown or white on the white apical spots; beset with short black setae. Forewing: membrane mostly hyaline except for a large transversal preapi- cal brown band, apical and posterior margins distal to the transversal band also brown, end of CuP+1A with a small curved brown mark, small brown infuscations at the base of crossveins around RA, and poststigmal area mostly white. Cubital fork basal to Rs origin; seven to eight presectoral crossveins; subcostal veinlets mostly simple, but the ones near the junction between Sc and RA forked; crossveins at the prefork area simple; prefork area about twice wider than posterior area. Hind wing membrane hyaline except by the apical third brown; brown area with three white marks: two at the hypostigmal cell level (one at the anterior margin and the other at the posterior margin and slightly larger) and the largest one on the apical region, comprising most of the anterior margin and brown indented on the posterior area. One presectoral crossvein. Prefork and posterior areas about the same width Legs (Fig. 3a, c): Tibia and femur about the same size and slightly longer than tarsi, (hind leg with tibia and femur about twice longer than tarsi). Coxa dark brown basally and pale apically, remaining segments mostly pale, except by small brown dots on the base of the setae, and the dark brown apex of the femur, tibia and T5; femur sometimes with dark brown mark near the base. All segments covered with black setae, except by the coxa with white setae. Tibial spurs long, passing T2 apex; tarsomeres length: T2, T3, and T4 about the same length, T1 as long as T2 and T3 combined, and T5 slightly longer than T1; claws shorter than half of T5 length. Foreleg: sense hair relatively short, only slightly longer than the regular setae; tibia with a patch of antennal cleaning setae on the apical region. Abdomen: All sclerites dark brown, except by the tergites 1 and 2 with a pale medial longitudinal line, and the base of tergites 4–6 pale. All sclerites covered by short black setae, and white setae on the pale areas of the tergites, and first and second segments covered by long white setae Female Terminalia (Fig. 4): Ectoproct rounded, set with thin black elongate setae dorsally and few thickened setae on the ventral area, in lateral view. Lateral gonapophyses smaller than ectoproct in lateral view, beset with thickened setae, which are slightly longer than the ones on the ectoproct. Tergite IX thin dorsally but broadening towards the ventral area in lateral view, set with short black setae and a row of thickened setae on the ventral margin. Sternite VIII short, in ventral view rectangular and with the distal margin straight. Pregenital plate small and conical. Posterior gonapophyses short with posterior margin rounded, covered with long black thickened setae, longer than the ones at the lateral gonapophyses. Anterior gonapophyses absent. Gonapophyseal plates elongate and parallel. Spermatheca with the distal half broader and darker, than proximal half. Male Terminalia: Unknown. Comments: Among the Brazilian species of Glenurus, G. incalis can be easily identified based on the presence of a large brown stripe on the forewing, the presence of a large indented white mark apically in the hind wing, and the presence of a single white mark at the posterior margin within the brown area in the hind wing (Fig. 2b) (G. peculiaris presents two white marks). Within the whole genus, G. incalis seems more similar to G. proi Navás, 1930 (known from Costa Rica, Honduras and Mexico) based on the indented white mark on the hind wing. However, they can be distinguished based on the pterothorax pleura color: entirely dark brown in G. incalis, but sharply divided in a dorsal dark brown line and a ventral pale brown line in G. proi (Stange 2002). As mentioned above, G. incalis is the least known species of the genus, and until now it was only recognized based on the female holotype from Peru. The holotype was collected in Chanchamayo in the Junín department in the Peruvian Amazon. Herein we report a new specimen also from Junín, at the Satipo valley, and another three specimens from the Brazilian Amazon, one from the Rondônia state, municipality of Vilhena, and the other two from the Amazonas state, with records for the municipalities of Manaus and Itacoatiara. Both places in the Amazonas states are located in the center of the Amazon basin, on the northern shore of the Amazon river. These new distribution records considerably expand the distribution of G. incalis, suggesting that the species could be widespread through- out the Amazon forest.

Redescription of Glenurus incalis Zootaxa 4858 (1) © 2020 Magnolia Press · 139 Figure 3. Glenurus incalis, a: head, frontal view; b: head and thorax, dorsal view; c: head and thorax, lateral view. Scale bars = 1 mm.

Examined specimens. Brazil: Amazonas: Itacoatiara, Fazenda Aruanã, AM 010, Km 215, 28–29.ix.1992, 21:00–22:00h, Motta C.S., Peralta F.A., Telos B.R., Hutchings R.S.G., Hamada N. (1♀ INPA); Manaus, 27.viii.1962, K. Lenko (1♀ INPA); Rondônia: Vilhena, 12º34’S–60º03’W, 615m, 11.ix.1999, Coleção Embrapa CPAC no 17484. (1♀ Embrapa-DF); Peru: Junín: Chanchamayo, N. Banks, Type 12030, MCZ-ENT 00012030 (♀ holotype MCZ); Satipo Valley, 600m, vii–viii.1987 (1♀ DZUP).

140 · Zootaxa 4858 (1) © 2020 Magnolia Press Machado Figure 4. Glenurus incalis female terminalia, a: terminalia, lateral view; b: terminalia, ventral view; c: spermatheca. e = ectoproct; gp = gonapophyseal plate; lg = lateral gonapophysis; pg = posterior gonapophysis; pp = pregenital plate; S = sternite; T = tergite.

Redescription of Glenurus incalis Zootaxa 4858 (1) © 2020 Magnolia Press · 141 Figure 5. South America map showing all the known records of Glenurus incalis, and the Brazilian records of Glenurus heteropteryx and Glenurus peculiaris.

1 Forewing with a transversal brown band at the apex (Fig. 2b–c) ...... 2 - Forewing without a transversal brown band at the apex (Fig. 2a)...... G. heteropteryx 2 Hind wing apex with two transparent marks at the posterior margin and the large transparent mark at the anterior margin not brown indented posteriorly (Fig. 2c)...... G. peculiaris - Hind wing apex with one transparent mark at the posterior margin and the large transparent mark at the anterior margin brown indented posteriorly (Fig. 2b)...... G. incalis

Acknowledgments

I would like to thank all the curators that made the specimens available to this study: Dr. Márcio Oliveira and Thiago Mahlmann (INPA); Dr. Eliana Cancello (MZUSP); Dr. Adolfo Calor and Leon Tavares (UFBA); Dr. Brígida Souza (UFLA); Dr. Amabílio Camargo (Embrapa-DF); Dr. Sinval Silveira Neto (Esalq); Dr. Cátia Patiu (MNRJ). I am thankful to Adrian Ardila-Camacho for providing the images of the holotype, to Dr. Fernando Vaz de Mello for al- lowing me to use his laboratory facilities, Vinicius Costa Silva for the help with the map, and Dr. Ricardo Kawada for the help with some images. I am grateful to two anonymous reviewers for their important suggestions. I am also grateful to the following funding agencies: Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq: 400237/2017-2) and Redes Regionais de Pesquisa em Biodiversidade e Biotecnologia No 79/2013 (grant 407.627/2013-8).

References

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