Zootaxa 4413 (2): 295–324 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2018 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4413.2.4 http://zoobank.org/urn:lsid:zoobank.org:pub:8F4A8473-CCE5-41AA-99A6-381CFB434586 New species and new distributional records of Neotropical (Insecta: )

ADRIAN ARDILA-CAMACHO1,4, ARLEY CALLE-TOBÓN2, MARTA WOLFF2 & LIONEL A. STANGE3 1Doctorado en Ciencias Biológicas, Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México, Mexico city—Mexico. Escuela de posgrado en Ciencias Biológicas, Departamento de Ciencias Biológicas, Universidad de los Andes, Bogotá—Colombia. 2Grupo de Entomología, Instituto de Biología, Universidad de Antioquia, Medellín—Colombia. E-mail: [email protected]; [email protected] 3Florida State Collection of , Gainesville, Florida, 32614-7100, U.S.A. 4Corresponding author. E-mail: [email protected]

Abstract

The Neotropical fauna of Mantispidae is currently composed of 106 species. We provide new distributional records of Mantispidae from Colombia and Panama. Three new species are described, one in Symphrasinae from Colombia, and two in Mantispinae from Colombia and Panama. Haematomantispa nubeculosa (Navás, 1933) and Leptomantispa axillaris (Navás, 1908) are reported from Colombia for the first time, the former being the first record of the genus in the country. New locality records for other species previously known from Colombia are also given. For Panama, we report Anchieta fasciatella (Westwood, 1867) and Trichoscelia iridella (Westwood, 1867) for the first time, the former is herein newly transferred from Plega to Anchieta. Three names confluens Navás, 1914, n. syn., Buyda apicata Navás, 1926, n. syn., and Mantispa neotropica Navás, 1933, n. syn., are here synonymized with Buyda phthisica (Gerstaecker, 1885). Updated keys for the genera of Mantispinae, and species of genera Trichoscelia, Buyda, and from Colombia are included. With this new information, the known species richness of Mantispidae from Colombia increases from 21 to 26, and from 16 to 19 species in Panama.

Key words: Mantispids, Neotropics, taxonomy, new records, new species

Introduction

The family Mantispidae (Neuroptera), commonly known as mantidflies, is a charismatic group of lacewings recognized by their morphological similarity to of the order Mantodea and members from other orders equipped with raptorial anterior legs (Frantsevich 1998; Kral 2013). Such resemblance is evidently product of evolutionary convergence (Aspöck & Aspöck 2007; Ohl 2011). As diagnostic traits, adults of this family present raptorial forelegs, with elongate coxa and robust femur equipped with small spines or tubercles, in addition to macrosetae and generally a prominent sub-basal spine. Also, the group presents an elongated and tubular prothorax behind the insertion of the forelegs and a ventrally fused pronotum forming a tubular structure (Aspöck & Mansell 1994; Ohl 2004; Liu et al. 2015). Mantispidae is composed of four subfamilies (Symphrasinae, Drepanicinae, Calomantispinae, and Mantispinae) which were found as monophyletic by Liu et al. (2015), but in a recent phylogenomic study the family was recovered as paraphyletic with two sister clades i.e. Rhachiberothidae + Symphrasinae and Berothidae + remaining Mantispidae (Winterton et al. 2017). Among the members of Neuroptera, Mantispidae is distinguished by the complexity of its development, since their immature stages suffer substantial morphological, physiological and behavioral changes through the different larval instars (New 1989; Hoffman & Brushwein 1992; Aspöck & Aspöck 2007). Although the mantidflies have been referred to as hypermetamorphic (Aspöck et al. 2001; Aspöck & Aspöck 2007), some authors indicate that the group does not have a true hypermetamorphosis as it is present in the meloid beetles (Redborg 1998; Monserrat 2014).

Accepted by A. Letardi: 13 Mar. 2018; published: 23 Apr. 2018 295 The larvae of Mantispidae also show specialized feeding habits since they predate or parasitize immature or adults of certain groups of arthropods, however within the different Mantispidae subfamilies there are species that can be generalists or specialists with regard of their host or prey association (Redborg 1998). Members of the subfamily Mantispinae are obligate predators on spider eggs, but interestingly some representatives can be ectoparasites of spider hemolymph temporarily while eggs are unavailable (Redborg 1998; Ohl 2011). On the other hand, the biology of the remainder subfamilies is largely unknown (Redborg 1998; Maia-Silva et al. 2013). Symphrasinae members are predators on larvae and pupae of holometabolous insects, including beetles, moths, flies, bees, and wasps (Hook et al. 2010). Among Calomantispinae members, immatures of Nolima pinal Rehn have been successfully reared in laboratory fed with various arthropods (MacLeod & Redborg 1982; Redborg 1998) whereas larvae of the Drepanicinae genus Theristria have been found associated with a web-building spider (Austin 1985) and the larvae of Ditaxis biseriata (Westwood) are likely edaphic (Dorey & Merrit 2017). The world fauna of Mantispidae is represented by just over 400 species distributed in all Biogeographic realms (Ohl 2004). The subfamily Symphrasinae is endemic to the New World, Drepanicinae has representatives in Asia, Australia, and the Neotropics, Calomantispinae is present in Central- and North America, as well as Australia, whereas Mantispinae is worldwide distributed (Ohl 2004; Liu et al. 2015). The taxonomy of the group is not completely understood, and there are vast areas around the world (e.g., South East Asia, New Guinea, Africa, and part of the Neotropical region), where the group is largely unknown (Ohl 2005; Snyman et al. 2012). Regarding the fauna of the New World, Hoffman (1992) presented a comprehensive revisionary work on Mantispidae in his doctoral dissertation. However, this important source has been only partially published (Hoffman 2002; Hoffman et al. 2017), thereby, many nomenclatural actions are not recognized as available (Ohl 2005). Meanwhile, some revisionary works were restricted to the study of the fauna from the Amazon basin (Penny 1982; Penny & Da Costa 1983). After the work of Hoffman (2002), a catalog of the family Mantispidae was published (Ohl 2004), and several important studies were carried out on the fauna from Mexico (Reynoso-Velasco & Contreras-Ramos 2008), Brazil (Machado & Rafael 2010), and Colombia (Ardila-Camacho & García 2015); describing new species, extending the known distribution for many others, and providing relevant illustrations and keys for species identification. Despite of these contributions, there are many gaps on the knowledge of Neotropical Mantispidae. For example, Symphrasinae and Drepanicinae need revisionary work, and regional taxonomic studies of the whole family are quite necessary (Ohl 2004, 2005). Until now, the Neotropical species richness of the family raises to 106 representatives. However, the taxonomic status of 19 species in the genus Mantispa Illiger, which does not occur in the New World (Hoffman 2002), need to be clarified (Ohl 2004; Machado & Rafael 2010; Ardila-Camacho & García 2015). In Colombia, 21 species of mantidflies have been reported, however, most of the extension of the country is still unexplored (Ardila-Camacho & García 2015). Moreover, the Panamanian fauna of Mantispidae is scarcely documented, and only 16 species have been reported so far (Hoffman 2002; Ohl 2004; Reynoso-Velasco & Contreras-Ramos 2008; Ardila-Camacho & García 2015; Hoffman et al. 2017). The main purpose of the present study is to describe three new species, one in the genus Trichoscelia Westwood and one in the genus Climaciella Enderlein from Colombia, plus one from the genus Buyda Navás from Colombia and Panama. Additionally, we present new distributional records and keys for selected genera of Neotropical Mantispidae.

Materials and methods

The specimens studied are housed in the Colección Entomológica de la Universidad de Antioquia, Medellín (CEUA), Museo Entomológico “Francisco Luis Gallego”, Universidad Nacional de Colombia, sede Medellín (MEFLG), Colección Entomológica de la Facultad de Agronomía, Universidad Nacional de Colombia, Bogotá (UNAB), Colección Taxonómica Nacional de Insectos “Luis María Murillo”, Corpoica, Mosquera, Cundinamarca (CTNI), Colección de Artrópodos de la Universidad Distrital Francisco José de Caldas, Bogotá (CAUD), Museo de Historia Natural de la Universidad de los Andes, Bogotá (ANDES-E), Florida State Collection of Artrhopods, Gainesville, Florida, USA (FSCA), Smithsonian Institution, Washington, D.C., USA (USNM), and University of Arkansas Museum, Fayetteville, Arkansas, USA (UAAM). Specimens genitalia were examined by dissecting the entire abdomen in the males and the last four abdominal segments in the females. These structures were cleared in a solution of 10% Potassium Hydroxide (KOH), and subsequently washed in distilled water, 10%

296 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. acetic acid and 70% ethyl alcohol. Finally, genitalia were stored in vials with glycerin. Examination of the external morphology was carried out using a Zeiss Stemi 2000 stereomicroscope. Photographs were taken with the same equipment, adapted with an AxioCam ERc 5s digital camera. Morphological terminology generally follows Hoffman (2002), except for the genitalia and wing venation, which are based on Lambkin (1986) and Aspöck & Aspöck (2008), and Breitkreuz et al. (2017), respectively. Acronyms of type depositories and international museums are the same used by Ohl (2004).

List of international museums cited in the text.

BMNH The Natural History Museum, London, Great Britain. CN Navás collection, partly destroyed. Some surviving remnants in the Museo de Zoología de Barcelona, Spain (MZBS). CZC collection F. de Zayas, Cuba. EMAU Ernst-Moritz-Arndt-University, Greifswald, Germany. INPA Coleção Sistematica da Entomologia, Manaus, Amazonas, Brazil. ISNB Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium. MACN Museo Argentino de Ciencias Naturales, Bueno Aires, Argentina. MNHN Museum National d'Histoire Naturelle, Paris, France. MZBS Museo de Zoología, Barcelona, Spain. MZPW Museum of the Institute of Zoology, Polish Academy of sciences, Warsaw, Poland. OUM University Museum, Oxford, Great Britain. ZMB Museum für Naturkunde, Berlin, Germany. ZMUH Zoologisches Institut und Zoologisches Museum, Hamburg, Germany.

Results

Taxonomy

Subfamily Symphrasinae

This subfamily is composed of 36 species grouped into three genera, Anchieta Navás, Plega Navás, and Trichoscelia Westwood (Penny 1982). The group has a distribution range encompassing from South United States to Argentina (Ohl 2004). Currently, Symphrasinae lacks a taxonomic revision and the generic limits are not well understood (Liu et al. 2015).

Genus Anchieta Navás, 1909

Type species: Anchieta nobilis Navás, 1909: 484 (=Anchieta fumosella (Westwood, 1867: 504)).

Anchieta fasciatella (Westwood, 1867), new combination Fig. 3a

Mantispa (Trichoscelia) fasciatella Westwood, 1867: 503. Holotype (or syntypes), female, Venezuela (OUM).

Specimens examined. New records: Panama: Canal Zone, Albrook Forest Site, 30.40 m, 25–26.VII.1968, black light trap, R. S. Hutton, (1 ♂—UAAM); same but 13‒14.VII.1967, Hutton & Llaurudo, (1 ♂—UAAM); same but 24‒25.VIII.1967, black light trap, Hutton & Llaurudo (2 ♂‒UAAM); same but 29.II‒1.III‒1968, R.S. Hutton, (1 ♂—UAAM); same but 3‒4.VIII.1967, Hutton & Llaurudo, (2 ♂—UAAM); Darién, Punta Patiño, 7.VIII.1952, F. Blanton (1 ♂—FSCA); Panamá, 7-10 km. north El Llano, 14‒22.V.1993, E. Giesberg (5 ♂—FSCA); Panamá Oeste, Barro Colorado Island, 18.IV.1874, Duckworth (1 ♂—USNM).

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 297 Distribution. Colombia (Bolívar, Magdalena, Santander; Ardila-Camacho & García 2015), Panama (Canal Zone, Panamá Oeste, Panamá, Darién). Comments. The genus Anchieta is until now composed of seven species distributed from Panama south to Brazil. Most of the species are known from the Brazilian Amazonia, although the genus presents few records (Penny 1982). The group is separated from other Symphrasinae members by the presence of a straight rarp2 (second anterior radial cell) of forewing. Another remarkable difference, is the mimetic pattern with several families of Hymenoptera, which seems to be a generalized feature of the group. For example, A. fumosella was referred to be a mimetic of Vespidae (Buys 2008), while A. fasciatella resembles wasps of the family Braconidae (Braconinae), which have repugnant abdominal glands (Ardila-Camacho & García 2015). The remainder species present a general color pattern and the hindtibia greatly swollen and flattened, resembling stingless bees of the tribe Meliponini (Penny 1982). Some species in this genus have lateral keel like projections on the abdominal tergites of both sexes. The pterostigma on the forewing is shorter than in Plega or Trichoscelia, and the trochanter of the forelegs has a dentiform process on the inner surface. Anchieta fasciatella was originally described from Venezuela. However, this record is now considered erroneous because the type locality, Santa Marta, is not located in Venezuela, but in Colombia. In fact, Ardila- Camacho & García (2015) reported this species as Plega fasciatella from several localities in Colombia. Moreover, in the revision by Penny (1982), this species was mentioned as occurring in Panama, however, no specific data were provided by the author. Here we present the first specific records of this species from Panama.

Genus Plega Navás, 1928

Type species: Symphrasis signata Hagen, 1877: 208.

Plega hagenella (Westwood, 1867)

Mantispa hagenella Westwood, 1867: 504. Holotype: male, “Amazonia”, unspecified locality (probably Brazil), (OUM). Mantispa cognatella Westwood, 1867: 506. Holotype: female, Venezuela (OUM).

Specimens examined. Colombia: Antioquia: San Jerónimo, Vda. Puente Blanco, fca. La Clarita, 23.XI.2005, light trap, A. Mercado (1 ♀–CEUA); Caucasia, Hda. La Candelaria, 880 m, 23.XI.2007, manual collection, GEUA (1 ♀–CEUA). New locality records. Colombia: Caquetá: Solano, Reserva Koreguaje, Jericó, Konsaya, La Raya stream, 0°32’37’’N–75°6’5’’W, 10 m, 28.IX.2016, Light trap, J. Hoyos (1 ♂—ANDES-E). Distribution. Bolivia, Brazil, Colombia (Antioquia, Chocó, Caquetá, Cundinamarca, Magdalena, Meta, Santander, Tolima), Costa Rica, Nicaragua, Panama, Trinidad, Venezuela (Ohl 2004; Ardila-Camacho & García 2015).

Genus Trichoscelia Westwood, 1852

Type species: Mantispa (Trichoscelia) fenella Westwood, 1852: 269.

This genus comprises small mantispids (forewing length ≤15 mm) distributed from Mexico to Argentina, perhaps with the noteworthy exception of Chile. With 15 representatives, it is the mantispid genus with the greatest diversity in the Neotropics so far (Ardila-Camacho & García 2015). The broadened distal article of labial palp and the lack of a sub-basal spine of fore femur are distinctive traits of this genus (Willman 1990). Penny (1982) and Penny & da Costa (1983) revised the species from the Amazon basin, where two species groups were recognized, the T. varia group with three species and the T. fenella group with six species. Three additional unplaced species were also listed by these authors. Then, Ohl (2004) transferred T. zikani to the genus Plega, while Ardila-Camacho & García (2015) transferred T. remipes to the genus Anchieta. In the revision of Colombian Mantispidae, Ardila- Camacho & García (2015) described three new species that were placed in the T. fenella group.

298 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. The biology of the group is poorly known, but the available literature cites some species of Polybiini (Vespidae) as host of their larvae (Penny 1982; Dejean & Canard 1990).

Trichoscelia iridella (Westwood, 1867) Fig. 3b

Mantispa (Trichoscelia) iridella Westwood, 1867: 503. Lectotype, male, “Amazonia”, no specific locality (BMNH). Mantispa (Trichoscelia) basella Westwood, 1867: 504. Holotype, male, “Amazonia”, no specific locality (OUM). Images studied.

Specimen examined. New record. Panama: Canal Zone: Albrook Forest Site, 30.4 m, black light trap, 28‒29.XI.1968, Hutton (1 ♂—UAAM). Diagnosis. Antenna piceous, wings completely hyaline, without marks; male genitalia with ectoproct ovoid in lateral view; sternite 9 spoon-shaped in lateral view, posterior margin produced into a short lobe, narrow and rounded in ventral view; gonocoxites 9 short, dorsally curved in lateral view, sinuous in ventral view, apically bifid; pseudopenis coiled, forming a single loop; gonarcus U-shaped. Distribution. Brazil, Ecuador, Panama (Canal Zone), Trinidad (Penny 1982). Comments. This species had been reported for Brazil, Ecuador and Trinidad (Penny 1982), here we register its presence in Panama. This small species of Trichoscelia is the most common one in the central Amazon region (Penny 1982). Adults are present all year round, but in highest numbers in May. Individuals have generally been collected in or near primary forest, using light traps (Penny 1982). Immature stages and other aspects of their biology remain unknown.

Trichoscelia umbrata Ardila-Camacho, n. sp. Figs. 1, 2

Type material. Holotype ♂, Colombia: Caquetá: Florencia, Macagual, 1°37’ N–5°36’ W, 300 m, 13.X.2014, C. García, light trap (1 ♂–UNAB). Holotype condition. Good, pinned, wings extended, abdomen cleared and stored in a glass microvial below the specimen. Diagnosis. Antenna with scape yellow at base and piceous at distal portion, pedicel and flagellum piceous, with 34 articles, slightly wider than long in frontal view. Forefemur yellow with brown suffusion on outer surface. Forewing with basal region of costal and subcostal spaces dark amber; membrane adjacent to apical and posterior margins with dark infuscations ranging from distal portion of CuP to wing apex. Hindwing with basal part of subcostal space dark amber; membrane adjacent to posterior and apical margins with dark infuscations. Male genitalia with ectoproct subtrapezoidal in lateral view, sternite 9 pentagonal in ventral view, posteromedially forming an acute angle. Gonocoxites 9 short and curved, equipped with four short spines, two apical and two preapical; gonarcus medially anterodorsally projected in lateral view. Description. Based on a single pinned male. Head. Labrum dark brown, clypeus yellow with a brown transverse stripe (Fig.1 d); maxillary and labial palpi amber covered with fine brown setae; frons with anterior half yellow (Fig.1 d), posterior part piceous; antenna with scape yellowish at basal half, piceous at distal half, pedicel piceous, flagellum piceous with 34 articles covered with short thick setae, basal 3/4 with flagellomeres wider than long in frontal view, the rest as long as wide (Figs. 1b, c). Vertex with a wide piceous stripe (Fig. 1c), area surrounding compound eyes with a mixture of yellow and piceous; compound eyes metallic silver; occiput yellow; postgena yellow. Thorax. Pronotum yellow, with longitudinal brown stripe dorsally, anterior margin brown with a row of long dark brown bristles; medial region with two raised regions covered with long dark brown bristles (Fig. 1c). Mesonotum yellow, with small brown areas on the scutum bearing long dark brown setae, remainder surface with yellow setae (Fig. 1c); metanotum yellow with diffuse brown suffusions on the scutum. Pteropleura completely yellow, covered with yellow setae (Fig. 1b). Legs. Foreleg with coxa and trochanter yellow, densely covered with yellow setae; femur yellow, except for

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 299 light yellow area medially on outer surface, entire surface densely covered with yellow setae (Fig. 1b); tibia yellow at inner surface, smoky yellow at outer surface; tarsi yellow. Mid- and hindlegs entirely yellow, densely covered with yellow setae (Fig. 1b). First tarsomere nearly as long as the remainder tarsomeres together; tarsal claws light brown; arolium present. Wings. Forewing 8 mm long, costal space narrow, with basal dark amber infuscation, 12 crossveins; subcostal space dark amber at base, two crossveins present, one basal and another at midlength; radial field with two crossveins; three veins leaving from rarp1 (first anterior radial cell), two from rarp2 (second anterior radial cell). Seven gradate crossveins present. Pterostigma six times as long as wide, piceous with a central yellow spot. Wing membrane hyaline, with alternate smoky and clear areas adjacent to posterior and apical margins, with seven dark areas in total; the first located between apex of CuP and first branch of CuA, remainder located between apical branches of longitudinal veins (Fig. 1a). Wing venation predominantly piceous, except proximal half of CuA, CuP, A1 and base of A2 that are yellow; all veins densely covered with setae of the same color as cuticule; posterior and apical margins alternating piceous and yellow. Jugal lobe and a2a31 (AP) cell dark brown at posterior margin (Fig. 1a). Hindwing 5.5 mm long, costal space reduced, narrow, with 4 crossveins; subcostal space dark amber basally; pterostigma elongated, curved, eight times as long as wide, piceous with a preapical yellow spot. Two or three longitudinal veins leaving from rarp1 (first anterior radial cell), rarp2 (second anterior radial cell) lacking longitudinal veins; five gradate crossveins. Wing membrane hyaline with alternate smoky and clear areas on posterior margin, four dark areas in total, the first located on first CuA branch, remainder dark areas between distal forks of longitudinal veins; apical margin smoky (Fig. 1a). Wing base yellow. Wing venation predominantly piceous, except C+Sc light brown and basal half of CuA, CuP and anal veins yellow (Fig. 1a); posterior wing margin alternating piceous and pale yellow, except basal 1/4 completely yellow; all veins densely covered with setae of the same color as cuticle. Male genitalia. Ectoproct subtrapezoidal in lateral view, with numerous long bristles, with a concavity posteroventrally, followed by a mushroom-like anteroventral projection (Fig. 2a). Sternite 9 pentagonal in ventral view, posteromedially forming an acute angle (Figs. 2e), entire surface with long bristles, spoon shaped in lateral view, with posterior margin at the level of anterior margin of ectoproct (Fig. 1a). Gonocoxites 9 short, arched in lateral and ventral view, anterior apex slightly widened with two rounded processes, posterior apex equipped with 2 apical and two preapical short spines (Figs. 2c, d, f). Gonarcus with lateral arms arched in lateral view, medial portion anterodorsally projected, medial lobe posteroventrally directed, rounded in ventral view (Fig. 2c). Mediuncus short forming an angle of 45° with longitudinal axis of abdomen. Pseudopenis coiled, extended beyond the remainder sclerites, forming two loops, apex posteriorly projected outside the insect body (Figs. 2a, c, f). Eymology. The specific epithet is derived from the Latin—umbratus—, meaning shaded, in reference to the smoky areas on the posterior and apical margins of the wing membrane of this species. Feminine adjective. Distribution. Colombia (Caquetá). Comments. Trichoscelia umbrata is similar to T. anae Penny, 1982, because both species share the wing color pattern, with alternate smoky and clear areas on membrane adjacent to posterior and apical margins, and the subcostal space basally infuscated (Figs. 1a; 3c). Forewing length is a common feature as well. In addition, both species have four spines of the male gonocoxite 9, however, antennal morphology and coloration, as well as body color pattern and morphology of the male genitalia are remarkably different. Trichoscelia anae was described by Penny (1982) from a single male collected in Rondonia, Brazil (Fig. 3c). In his revision of the Brazilian Mantispidae, Penny & da Costa (1983) proposed a classification to distinguish species of Trichoscelia based on morphology of the antennal flagellomeres, in a similar way as Parker & Stange (1965) classified species of Plega. The T. fenella group includes species with antennal flagellomeres wider than long, while the varia group comprises species with antennal flagellomeres as long as wide. Along with T. varia, Penny classified T. anae within the T. varia group, and the rest of species were classified in the T. fenella group. Among distinctive traits of T. anae, the flagellomeres 8‒12 from the apex are pale yellow, character that separates these two species with the naked eye, since T. umbrata has antennae completely piceous. In addition, considering the morphology of the antennal flagellomeres, T. umbrata belongs to the T. fenella group. Body color pattern also differs. For instance, T. anae has pterothorax yellow with piceous spots in different parts of the sclerites, making a mottled pattern (Fig. 3c). In contrast, T. umbrata has the pterothorax nearly entirely yellow, with only a few brown suffusions on the thoracic nota. The forelegs are yellow with piceous spots on femur and tibia in T. anae, but in T. umbrata those are predominantly yellow. Mid- and hindlegs of T. anae are mottled as well (Fig. 3c), while in T. umbrata they are completely yellow. Male genitalia, are well distinguished between both species. The ectoproct of T. anae is

300 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. rounded, while in T. umbrata is subtrapezoidal; sternite 9 is posteromedially rounded in T. anae, but in T. umbrata is pentagonal. The shape, size and arrangement of the apical spines of gonocoxites 9 are distinct between these species. A comprehensive comparison of morphological differences between T. anae and T. umbrata is presented in Table 1. Considering such background of morphological differences, and the distant geographical distribution, we prefer to treat these species as separate rather than considering them as a single species.

TABLE 1. Morphological comparisons between T. anae Penny and T. umbrata n. sp. Character Trichoscelia anae Penny Trichoscelia umbrata n. sp. Antennal flagellomeres as long as wide slightly longer than wide morphology Antennae color pattern 5 preapical flagellomeres light yellow completely piceous Number of antennal articles 32 34 Pronotum anterior margin and two median raised spots longitudinal stripe and anterior margin piceous piceous Pterothorax color yellow with piceous spots almost completely yellow Foreleg color coxa and trochanter yellow with piceous coxa and trochanter yellow with some diffuse spots; femur inner surface with basal brown light brown areas; femur almost completely spot, distal half of surface adjacent to the yellow with a light brown area on outer surface inner row of setae brown Tibia color yellow with piceous spots dorsally yellow on inner surface, smoky on outer surface Mid- and hindlegs color mid and hind femur with brown spots basal- completely yellow and apically; midtibia brown basally, hindtibia piceous with a yellow band medially FW costal space distinctively widened at midlength, 15 slightly widened at midlenght, 12 crossveins crossveins present, entirely hyaline present, basally with fuscous spot FW gradate crossveins six seven FW veins color venation mottled, alternating piceous and principal veins mainly piceous with a few yellow regions light brown; basal half of CuA yellow, CuP yellow except apex piceous; A1 yellow and A2 mottled; posterior and apical wing margin alternating yellow and piceous FW longitudinal veins rarp1:2, rarp2: 2 rarp1:3, rarp2: 2 leaving from anterior radial cells (i.e. radial formula) HW costal space slightly widened, 3 crossveins narrow, 2 crossveins HW subcostal space piceous with a small fuscous spot with a wide basal fuscous spot HW veins color venation mottled, alternating piceous and venation nearly entirely piceous, C+Sc smoky yellow yellow; wing base yellow, basal half of CuA yellow, CuP yellow, except piceous apex; anal veins yellow; posterior wing margin with basal 1/4 yellow, the rest alternating piceous and yellow, apical margin piceous HW radial formula rarp1:3, rarp2: 1 rarp1:2, rarp2: 0 Ectoproct rounded in lateral view subtrapezoidal in lateral view Sternite 9 laterally spoon shaped, posteromedially laterally spoon shaped, pentagonal, rounded, with apex extending beyond of the posteromedially forming an acute angle, level of the posterior margin of ectoproct extending to posterior margin of ectoproct Mediuncus long, subparallel with the longitudinal axis short, directed at 45° from the longitudinal axis of the abdomen of the abdomen ...... continued on the next page

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 301 TABLE 1. (Continued) Character Trichoscelia anae Penny Trichoscelia umbrata n. sp. Pseudopenis long, forming a single loop, apex apparently long, coiled, forming two loops behind the not posteriorly projected outside the insect mediuncus, apex posteriorly projected outside body the insect body Gonocoxites 9 long in lateral view, with apex distinctively short in lateral view, slightly arched in ventral curved dorsad in ventral view, sinuous in view, with two short apical and two preapical ventral view, apex curved laterally, with four spines long apical spines

In a recent work by Liu et al. (2015), the genitalia of T. anae was illustrated (Figures 11F, G) and the figures presented by the authors were more similar to the holotype of T. umbrata, possessing only few minor differences in the shape of the ectoproct and sternite 9.

FIGURE 1. Trichoscelia umbrata n. sp.: a, wing venation; b, head and thorax, lateral view; c, head and thorax, dorsal view; d, head, frontal view.

302 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. FIGURE 2. Trichoscelia umbrata n. sp.: a, male genitalia, lateral view; b, same, ventral view; c, male genitalia, lateral view; d, apex of gonocoxite 9, lateral view (above), ventral view (below); e, sternite 9; f, male genitalia, ventral view. Abbreviations: g, gonarcus; gcx9, gonocoxite 9; hpm, hipomere; med, mediuncus; psp, pseudopenis. Scale bars: 0.1 mm.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 303 Key to Colombian species of Trichoscelia (after Ardila-Camacho & García 2015, modified)

1. FW pterostigma mostly piceous without yellow spots (Ardila-Camacho & García 2015, Figs. 14a, b); gonocoxite 9 with one long preapical spine (Ardila-Camacho & García 2015, Fig. 15d) ...... T. karijona Ardila-Camacho, 2015 - FW pterostigma brown with a central yellow spot; gonocoxite 9 with four to six apical spines ...... 2 2. Posterior and apical margins of forewing with a series of amber infuscations (Fig.1); male gonocoxite 9 with four apical spines (Fig. 2d)...... T. umbrata n. sp. - Forewing nearly entirely hyaline; gonocoxite 9 with five or six apical spines...... 3 3. Area between CuA and posterior wing margin of both wings with membrane and venation light yellow (Ardila-Camacho & García 2015, Figs. 12a, b); forefemur completely yellow; male gonocoxite 9 with five apical spines (Ardila-Camacho & García 2015, Fig. 13c) ...... T. gorgonensis Ardila-Camacho, 2015 - Area between CuA and posterior wing margin of both wings with venation alternating brown and yellow and membrane hya- line; forefemur yellow but brown infuscated dorsolaterally (Ardila-Camacho & García 2015, Figs. 9b, c); male gonocoxite with six apical spines...... T. andina Ardila-Camacho, 2015

Subfamily Mantispinae

Currently the New World fauna of this subfamily is composed of approximately 58 species grouped into ten genera (Buyda Navás, Climaciella Enderlein, Dicromantispa Hoffman, Entanoneura Enderlein, Haematomantispa Hoffman, Leptomantispa Hoffman, Mantispa Illiger, Paramantispa Williner & Kormilev, Xeromantispa Hoffman, and Zeugomantispa Hoffman). In the Americas, this group is found from Canada to Argentina but it is absent in Chile (Ohl 2004).

Genus Buyda Navás, 1926

Type species: Buyda apicata Navás, 1926: 87.

Before this study, the genus Buyda was represented in the Neotropics by two species, B. phthisica (Gerstaecker) and B. apicata Navás. The second species is herein synonymized with the former, as proposed in the revisionary work by Hoffman (1992), being B. phthisica the only valid species of this genus until now. The geographical distribution of this species ranges from Honduras to southern Argentina (Hoffman 2002). Here, we describe a new species for this genus, which was previously described from Panama in Hoffman’s Ph.D. dissertation of 1992 but was not formally published by the author. This new species presents a body color pattern, genital morphology and arrangement of the abdominal pregenital apparatus remarkably different from B. phthisica. Ardila-Camacho & García (2015) reported this species as inquirenda from the departments of Antioquia and Sucre based on two female specimens. According to Ardila-Camacho & García (2015), the examination of male specimens could clarify the identity of these specimens, either as specimens pertaining to B. phthisica or as the species described as new in the Hoffman´s dissertation, given the morphological similitude of female specimens. In the present study, we describe this species with consent of K.M. Hoffman (pers. comm.). The species description is based on the work by Hoffman (1992) and our own examination of a series of specimens housed in CEUA and ANDES-E.

Buyda immaculata Ardila-Camacho, Calle-Tobón & Wolff, n. sp. Figs. 3d, 4–6

Type material. Holotype ♂, Colombia: Antioquia: Puerto Berrío, Alto de las Águilas, Hacienda Manaos, 6°27’10.09’’ N−74°36’13.74’’ W, 440 m, 06.VII~07.VII.2013, Wolff-INPA, white light trap, 18:00−19:00 [Forewing length, 10.1 mm; Hindwing length, 9.4 mm] (CEUA). Paratypes: Colombia: Antioquia: Puerto Berrío, Alto de las Águilas, Hacienda Manaos, 6°27’10.09’’ N−74°36’13.74’’ W, 440 m, 06–07.VII.2013, Wolff- INPA, light trap, 18:00–19:00 (1 ♀−CEUA); Sucre: Colosó, Montes de María, II.2012, J. Noriega, light trap in forest (2 ♀—ANDES-E).

304 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. Holotype condition: good, pinned, with right wings extended, abdomen dissected and stored in a microvial with glycerin below the specimen. Diagnosis. Body coloration pattern consisting in a mixture of dark brown and light yellow, wing membrane entirely hyaline. Male abdomen bearing two patches of 30−55 circular pores laterally, on each side close to the anterior margin of tergite 5. Intersegmental membrane between tergites 5 and 6 invaginated, forming a bilobed pocket extended anteriorly 1/5 the length of the tergite 5. Ectoproct ovoid, rounded at apex. Sternite 9 subpentagonal in ventral view, with apex broadly rounded. Hypomeres elongated, shorter than pseudopenis, apex slightly clavate. Female terminalia with sternite 8 as long medially as laterally. Spermatheca with distal section slightly wider than medial section. Description. Based on a pinned male and female, two females preserved in alcohol and the description by Hoffman (1992). Head. Labrum dark brown, clypeus brown medially; mandibles dark amber; maxillary and labial palpi yellow to yellowish brown. Frons with a brown V-shaped inverted mark, medially with a pentagonal region light yellow in color (Fig. 4c). Scape light yellow anteriorly, brown posteriorly, pedicele brown, flagellum with 28−33 articles covered with short and thick setae. Vertex mainly brown with yellow spots near compound eyes (Fig. 4c); postgena yellow (Fig. 4d). Compound eyes metallic brown. Thorax. Prothorax 3.5−5 mm long, straight in lateral view (Fig. 4b); pronotum light yellow or tan, with a brown longitudinal stripe at midline (Fig. 4d), prozona with two pairs of lateral brown spots, anterior margin brown, maculae brown (Fig. 4d); medial region laterally with two elongate brown spots (Fig. 2c); posterior portion with a brown spot. Mesonotum light yellow or tan to light brown, with two pairs of longitudinal brown stripes, scutellum with a V-shaped brown mark (Fig. 4d). Metanotum light brown with two lateral dark brown spots and light yellow at middle; scutellum with a V-shaped brown mark (Fig. 4d). Pteropleura mottled, sclerites mainly brown with some peripheral yellow regions (Fig. 4b). Legs. Foreleg with coxa light brown, dorsal- and ventrally with longitudinal brown stripes; trochanter varying from light to dark brown. Femur 4–5 mm long, light or dark brown at inner surface (Fig. 4f); sub-basal spine light to dark brown; outer surface yellow to light brown, with brown spots at base and apex, medial region with a brown spot adjacent to row of minor spines; with 30–43 minor spines, basally yellow and brown at apex, (Figs. 4b, e). Tibia dorsally light to dark brown, ventrally with dark brown stripes. Tarsi brown. All segments covered with fine dark brown or yellow setae. Mid- and hindleg with coxa and trochanter light to dark brown, femur and tibia yellow to light brown, femur dark brown at base and apex, tibia basally dark brown. Tarsi light brown; tarsal claws brown with four apical teeth; arolium present. All segments covered with fine light or dark brown or yellow setae. Wings. Forewing 10.1–13 mm long, membrane hyaline, venation brown; costal space with six or seven crossveins. Pterostigma brown (Fig. 4a). Subcostal space hyaline to slightly infuscate, with four or five crossveins. RP with 5 or 6 branches, one branch arising from rarp1 (first radial cell), two or three leaving from rarp2 and rarp3 (second and third anterior radial cells). a2a3-1 with asperous region dark brown. A1 and base of A2 yellowish brown basally. Hindwing 9.4–12 mm long, similar to forewing, membrane hyaline. Pterostigma brown. Principal veins brown, except base of Cu and anal veins that are yellow or light brown (Fig. 4a). Costal space with six or seven crossveins. Abdomen. Tergites light brown, with brown stripe located on each sclerite on the posterior region. Sternites light brown, the first five with yellow margins, sternites of abdominal segments 6 and 7 dark brown. Pleura yellow to dark brown. Male pregenital abdominal apparatus: tergite 5 with anterolateral oval patches of 30–55 circular pores. Intersegmental membrane between the abdominal segments 5 and 6 invaginated into a bilobed pocket extending anteriorly 1/5 the length of tergite 5 (Figs. 5a, c, d). Male genitalia. Ectoproct brown, ovoid, with apex rounded, covered with long fine setae (Figs. 5a, b). Ventromedial lobe unsclerotized, directed medially, rounded at apex, with 20–30 short and thick setae. Sternite 9 subpentagonal, apex broadly rounded in ventral view (Fig. 5i). Gonarcus median lobe wider than long, rounded at apex (Fig. 1h). Mediuncus widened at base in ventral view, apex forked (Fig. 1g). Gonocoxites slightly wider distally than basally, apex rounded and bent dorsolaterally. Pseudopenal membrane short. Hypomeres elongate, curved, shorter than pseudopenis, apex slightly knobbed (Fig. 5g). Pseudopenis sclerotized, curved dorsad in lateral view, pointed at apex, spine-like (Figs. 5g). Female. Body color pattern, similar to that of male. Pronotum length, 4–5 mm; forewing length 11–13 mm, hindwing length, 10–12 mm. Ectoproct ellipsoid (Figs. 6a, b), brown, covered with long yellow setae. Gonocoxite

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 305 9 ellipsoid (Figs. 6a, b), brown. Sternite 8 with a transverse brown stripe, posterior margin yellow to light brown, as long medially as laterally in ventral view (Figs. 6c, d). Spermatheca with medial section forming 1.5 coils wider than proximal section, distal section C-shaped, slightly wider than medial section (Figs. 6e, f). Fertilization canal duct shallowly curved at base, slightly sinuous distally (Figs. 6e, f). Etymology. As a tribute to Kevin M. Hoffman, we decided to name this species using the same name he originally utilized in his dissertation, immaculata, meaning unspotted, which refers to the lack of an apical infuscation on the wings, a distinctive trait of this species. Feminine adjective. Distribution. Colombia (Antioquia, Sucre), Panama (Hoffman 1992). Comments. As in B. phthisica, this species has a camouflage-like coloration pattern, consisting in a mixture of yellow and brown, but the absence of green color on the body, and the lack of an apical infuscation on the wings easily separate both species. The arrangement of the pregenital abdominal apparatus is another important character to distinguish these species. In B. phthisica, pores on the tergite 5 are lacking, while there are two anterolateral patches of pores in B. immaculata. Male genitalia characters (such as morphology of ectoproct, gonocoxites, hypomeres, gornarcal membrane and mediuncus) and female genitalia traits (such as sternite 8 and spermatheca) are different as well.

FIGURE 3. Habitus of Mantispidae spp.: a, Anchieta fasciatella (Westwood), male (photo by David Bowles); b, Paralectotype of Mantispa (Trichoscelia) iridella Westwood, female (photo by Amoret Spooner, OUM, Oxford); c, Holotype of Trichoscelia anae Penny, male (photo by Thiago Mahlmann, INPA, Manaus); d, Buyda immaculata n. sp.; e, Climaciella obtusa Hoffman, male (photo by Guillermo Guarin-Candamil, MEFLG, Medellín); f, Leptomantispa axillaris (Navás), male.

306 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. FIGURE 4. Buyda immaculata n. sp.: a, fore- and hindwing; b, head and thorax, lateral view; c, head, frontal view; d, head and thorax, dorsal view; e, forefemur, outer surface; f, same, inner surface. Abbreviations: mac, macula.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 307 FIGURE 5. Buyda immaculata n. sp.: a, cleared abdomen of male, lateral; b, male genitalia, dorsal view; c, abdominal segments 3‒6, dorsal view; d, same, lateral view; e, male genitalia, dorsal view; f, same, lateral view; g, male genitalia, ventral view; h, gonarcus, ventral view; i, sternite 9, ventral view. Abbreviations: g, gonarcus; gcx9, gonocoxite 9; hpm, hipomere; Mbn, intertergal membrane; med, mediuncus; po, pores; psp, pseudopenis; t3‒t6, third to sixth abdominal tergites; vml, ventromedial lobe. Scale bars: 0.1 mm, except for c.

308 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. FIGURE 6. Buyda immaculata n. sp.: a, b, female genitalia, lateral; c, d, same, ventral view; e, f, spermatheca, lateral view. Abbreviations: cap, capsule; ds, distal section of the spermatheca; ect, ectoproct; fc, fertilization canal; gcx9, gonocoxite 9; ms, medial section of the spermatheca; s7, s8, sternites of abdominal segments 7 and 8; sp, spermatheca; t8, t9, tergites of abdominal segments 8 and 9. Scale bars: 0.1 mm.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 309 Buyda phthisica (Gerstaecker, 1885)

Mantispa phthisica Gerstaecker, 1885 [“1884”]: 35. Holotype: female, Brazil (EMAU). Entanoneura phthisica (Gerstaecker, 1885), Handschin 1960: 208. Mantispa confluens Navás, 1914: 19. Holotype: sex not indicated, Panama (BMNH); New synonym. Buyda apicata Navás, 1926: 87. Holotype: female, type locality not indicated (MZBS); New synonym. Mantispa neotropica Navás, 1933: 309. Holotype: female, French Guiana (MNHN); New synonym.

New locality records. Colombia: Casanare: Aguaclara, “Station service”, 630 m, 13.IX.2015, 4°59’56’’ N– 72°47’13’’ W, C. Gonzalez, light trap (1 ♂—ANDES-E). Distribution. Argentina, Brazil, Colombia (Casanare, Meta, Valle del Cauca), Costa Rica, French Guiana, Nicaragua, Panama, Peru, Surinam, Uruguay (Hoffman 1992). Comments. The genus Buyda was described by Navás (1926) based on the species B. apicata, whose type specimen lacked specific locality data. The author mentions that the species was probably from Central Africa due to similarities with Pseudoclimaciella tropica (Westwood, 1852), and other African species (Navás 1926; Ohl 2004). However, in the list by Monserrat (1985) of the Mecoptera and Neuroptera from the Navás collection (CN) deposited in MZBS, it is indicated that the type specimen comes from Costa Rica (Ohl 2004). Then, Hoffman (1992) emphasized that this genus is an exclusive Neotropical element (Ohl 2004). In the original description of B. apicata, Navás (1926) illustrated the wing apex of the right wing, where the apical infuscation distinctive of B. phthisica is evident (Hoffman 2002). Furthermore, in a previous work, Navás (1914) described Mantispa confluens from Central America, specifically from Panama. In that publication, the author illustrated the wing apex and pronotum, which are undoubtedly diagnostic characters of B. phthisica. Additionally, after the description of B. apicata, Navás (1933) proposed a Neotropical species whose diagnostic traits (e.g., apical infuscation of wings and pronotum color pattern) perfectly match with those of B. phthisica. The species in question is Mantispa neotropica, and it was described based on a single female specimen deposited in the MNHN. Even, the specimen was labeled as Entanoneura phthisica (Gerstaecker, 1885) by R.G. Beard in 1968 (Tauber et al. 2017). In this work we synonymize these species with B. phthisica as it was already suggested by Hoffman (1992).

Key to species of Buyda (after Hoffman (1992), modified)

1. Wings apex hyaline (Fig. 4a); male abdominal tergite 5 with anterolateral patches of 30–55 pores (Fig. 5c); ectoproct rounded distal to ventromedial lobe (Fig. 5e); hypomeres shorter than pseudopenis (Fig. 5g); female sternite 8 as long medially as later- ally (Fig. 6c); spermatheca with distal section slightly wider than medial section (Fig. 6e) ...... B. immaculata n. sp. - Wings apex infuscated (Machado & Rafael 2010, Fig. 3); male tergite 5 without pores; ectoproct conical distal to ventromedial lobe; hypomeres nearly as long as pseudopenis (Machado & Rafael 2010, Fig. 4e, f); female sternite 8 nearly 1.5 times as long medially as laterally; spermatheca with distal section narrower than medial section (Machado & Rafael 2010, Fig. 4i) ...... B. phthisica (Gerstaecker, 1885)

Genus Climaciella Enderlein, 1910

Type species: Mantispa brunnea Say in Keating, 1824: 309.

The genus Climaciella comprises conspicuous and colorful mantispids. Little is known about this group whose species have a remarkable pattern of Batesian mimicry with polistine wasps (Opler 1981; Hoffman et al. 2017). However, some behavioral and ecological aspects of the Central and North American species Climaciella brunnea Say have been studied (Opler 1981). The rest of the Climaciella species inhabit across of the Neotropical region and for these species their respective models have not been identified. Immature stages are likely associated with spiders of the families Ctenidae and Lyscosidae, and the larvae of C. brunnea have been reported to be spider boarders (Redborg 1998). The genus is composed by eight species with a distribution ranging from southern Canada to northern Argentina, including the Greater Antilles (Hoffman 2002; Ohl 2004). Four species were recorded for Colombia. Climaciella amapaensis from Risaralda, C. porosa from the lowlands of the Anchicayá basin in the department of Valle del Cauca, C. semihyalina from the Colombian Amazonia and C. obtusa from the

310 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. Colombian Pacific coast and Caribbean region (Ardila-Camacho & García 2015). The genus is easily distinguished by the flagellum possessing flagellomeres three times wider than long at midlength in anterior view, pronotum straight or angled at midlength in lateral view, pterothorax with mesoscutal furrow not apparent, and anterior half of wings light to dark amber (Hoffman 1992; Machado & Rafael 2010).

Climaciella rafaeli Calle-Tobón, Ardila-Camacho & Wolff, n. sp. Figs. 7, 8

Type material. Holotype ♂, Colombia: Antioquia: Puerto Berrío, Alto de las Águilas, Hacienda Manaos, 6°27’10.09’’ N–74°36’13.74’’ W, 440 m, 6–7.VII.2013, Wolff-INPA, light trap, 18:00–19:00 [forewing length, 14.1 mm; hindwing length, 12.3 mm] (CEUA). Paratypes. None. Holotype condition. Good, pinned, apex of left antenna lacking, right wings extended, abdomen dissected and cleared, stored in microvial with glycerin below the specimen. Diagnosis. Body nearly completely black; prothorax ventrally reddish brown, bent ventrad at midlength in lateral view; pleural sclerites of pterothorax reddish brown at middle, black at the periphery. Wings with anterior half light amber, remainder hyaline. Foreleg with apex of coxa and trochanter reddish brown, femur black with a light reddish-brown stripe at the level of major spine, on inner and outer surfaces. Abdominal tergites 4 and 5 with anterolateral patches of pores, each composed of 10‒15 semicircular pores arranged in two subparallel irregular rows, which merge near dorsal midline on each side; inside of each patch an elongate scar is present. Pseudopenis slightly longer than pesudopenal membrane, spine-like. Description. Based on a single pinned male Head. Nearly entirely black (Fig. 7c); labrum mainly black, dark reddish brown at margins; Mandibles very dark amber; maxillary palpi black, posterior half of last palpomere reddish brown; labial palpi black, dark reddish brown on the junctions. Postgena light brown (Fig. 7d). Scape and pedicel dark reddish brown, flagellum black, composed of 29 articles, nearly two times as wide as long, all densely covered with short and thick black setae. Compound eyes metallic silver. Thorax. Prothorax 3.3 mm long, striated, bent ventrad at midlength in lateral view, anteriorly expanded in dorsal view, black, ventrally reddish brown (Figs. 7b, d). Meso- and metanotum black (Fig. 7d), pteropleura with sclerites reddish brown at middle and black at the periphery (Fig. 7b); entire surface covered with fine and short clear setae. Legs. Foreleg with coxa predominantly black, apex light brown, entire surface covered with fine and short clear setae; trochanter reddish brown; femur 4.5 mm long, mainly black with a transverse light reddish brown stripe at the level of major spine on inner and outer surfaces, and area adjacent to row of minor spines (Figs. 7e, f); tips of minor spines amber, major spine dark reddish brown; with 24 minor spines; entire surface covered with fine and short setae of the same color as cuticle. Tibia dark reddish brown on inner surface, black on outer surface, completely covered with fine and short dark setae. First tarsomere amber, light brown at apex, remainder tarsomeres together nearly as long as the first one, light brown with fine and short setae of the same color as cuticle. Mid- and hindleg black, tarsi dark reddish brown, tarsal claws amber with three apical teeth; arolium present. All segments densely covered with fine and short light-yellow setae. Wings. Forewing 14.1 mm long, costal field with eight crossveins; subcostal space with five crossveins. Anterior 1/2 light amber, venation brown (Fig. 7a); pterostigma light brown with minute setae. 10 branches arising from RP, four veins arising from rarp1 (first anterior radial cell), and three from rarp2 and rarp3 (second and third anterior radial cells). Hindwing similar to forewing, 12.3 mm long; costal space with 10 crossveins. Eight branches arising from RP, two from rarp1 (first anterior radial cell), and three from rarp2 and rarp3 (second and third anterior radial cells). Abdomen. Nearly completely black, covered with fine and short light-yellow setae. Male pregenital abdominal apparatus: tergites 4 and 5 bearing two anterolateral patches of semicircular pores arranged in two subparallel irregular rows, which merge near dorsal midline on each side, with 28 pores on tergite 4, and 22 pores on tergite 5; inside of each patch an elongate scar present. Intertergal membrane between tergites 5 and 6 invaginated, slightly bilobed.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 311 Male genitalia. Ectoproct elongate, ovoid in lateral view covered with long setae, ventromedial lobe lightly sclerotized, dorsoventrally flattened, bearing 35‒40 short and thick spinous setae (Figs. 8a, d). Sternite 9 pentagonal, posteromedially produced into a bluntly rounded lobe (Fig. 8b). Gonarcus median lobe short and pointed (Fig. 8h), nearly as long as mid-length of pseudopenis. Gonocoxites 9 slightly shorter than mediuncus, somewhat straight, subapically widened (Fig. 8f). Mediuncus basally rounded, forked at apex, with attenuated projections (Fig. 8f). Hypomeres present as a pair of small lateral elliptical sclerites. Pseudopenal membrane broadly triangular, pseudopenis spine-like, as long as pseudopenal membrane (Figs. 8f). Female. Unknown. Distribution. Colombia (Antioquia). Etymology. Named after Jose Albertino Rafael, Brazilian Entomologist who collected the holotype together with one of the authors (MW). Comments. Species very similar to C. obtusa, since both have resembling body color pattern and prothorax shape. They can be separated by the wing color pattern, C. rafaeli has the anterior half of both wings light amber, while C. obtusa presents anterior 3/4 of both wings dark amber, with metallic blue appearance when observed from certain angles. The tip of the antennae is yellow in C. obtusa, but it is black in the new species. Another important difference is the number of pores and morphology of the abdominal pregenital apparatus; C. rafaeli has a few pores irregularly arranged with a transverse elongate scar inside each patch of pores. In contrast, C. obtusa has numerous pores arranged as linear rows, but the scars are absent. The gonocoxites are relatively straight in the apical portion in the new species, but in C. obtusa the gonocoxite apex is laterally projected, forming a rounded lobe. The pseudopenis of C. rafaeli is spine-like, while C. obtusa has the pseudopenis apex truncate to broadly rounded. The shape of the apical fork of the mediuncus is markedly different as well.

Climaciella obtusa Hoffman, 2002 Fig. 3e

Climaciella obtusa Hoffman, 2002: 254, male, female. Holotype: male, Ecuador (FSCA)

Specimen examined. Colombia: Valle del Cauca: El Darien, 9.V.2017, no collector, mist net for birds (1♂— MEFLG). Diagnosis. Body almost totally black, posterior margin of eyes orange, pronotum very dark reddish brown ventrally. Foreleg with apex of coxa, trochanter and area surrounding the sub-basal spine at outer and inner surfaces orange-reddish. Prothorax bent ventrad at mid-length in lateral view. Wings with anterior 3/4 dark amber, remainder membrane hyaline (Fig. 3e). Male tergites 5 and 6 with anterolateral patches of pores, each one composed by 18–36 circular pores; intersegmental membrane between these segments expanded into a bilobed pocket, extended 1/3 the length of the tergite 5. Male genitalia with ventromedial lobe of ectoproct flattened, bearing 50–65 short and thick setae. Sternite 9 pentagonal in ventral view, scoop-like in lateral view, posteromedially produced into a short and rounded lobe. Gonarcus median lobe spine-like. Hypomeres present as paired ellipsoid lateral sclerites. Pseudopenis spine-like in posterior view, slightly longer than pseudopenal membrane, apex broadly rounded to truncate, with basal half or more as wide as base. Pseudopenal membrane broadly triangular. Distribution. Colombia (Magdalena, Valle del Cauca), Costa Rica, Ecuador, Panama (Hoffman 2002). Comments. This mantispid species can be found in lowland forests of Central America and Northern South America from 400 to 1450 m.a.s.l. (Hoffman 2002; Ardila-Camacho & García 2015). Adults have been collected in February, May to August and October (Hoffman 2002; Ardila-Camacho & García 2015). The immature stages and spider hosts are unknown. In the same way as C. semihyalina and C. brunnea (“synoeca” morph) (Opler 1981), C. obtusa has a Batesian mimicry pattern resembling some vespid wasp species of the genera Polybia Lepeletier and Synoeca de Saussure, although the possible species model have not been identified (Hoffman 2002; Ardila- Camacho & García 2015).

312 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. Climaciella semihyalina (Le Peletier de Saint-Fargeau & Audinet-Serville, 1825)

Mantispa semihyalina Le Peletier de Saint-Fargeau & Audinet-Serville, 1825: 270. Holotype: sex unknown, Brazil (MNHN). Mantispa chalybea Erichson, 1839: 160. Syntypes, sex unknown, Brazil, Suriname (ZMB, MCZ). Nobrega tinctus Navás, 1914: 233. Holotype, sex unknown, Brazil (BMNH).

Specimen examined. Colombia: Putumayo: Mocoa, vda. Callyaco, Jardín botánico, 1°7’6.3’’N–76°37’58.7’’W, 548 m, 2.III.2016, entomological net, J. Mora (1♂–UNAB). Distribution. Argentina, Bolivia, Brazil, Colombia (Putumayo), Ecuador, French Guiana, Paraguay, Peru, Suriname, Uruguay (Hoffman 1992). Comments. This species is distributed from the Amazon basin south to Argentina, but has an erroneous distribution range as north as to Mexico (Hoffman 1992; Reynoso-Velasco & Contreras-Ramos 2008).

Key to Colombian species of Climaciella (after Ardila-Camacho & García 2015, modified)

1. Body predominantly black; prothorax bent ventrad at midlength in lateral view (Ardila-Camacho & García 2015, Fig. 21a)...... 2 - Body predominantly dark reddish brown or with a mixture of black and yellow; prothorax straight in lateral view (Ardila- Camacho & García 2015, Fig. 17a)...... 4 2. Wings with anterior 3/4 dark amber, remainder membrane hyaline (Fig. 3e) ...... C. obtusa Hoffman, 2002 - Wings with anterior 1/2 amber ...... 3 3. Wings with posterior 1/2 completely hyaline (Fig. 7a); forefemur with a transverse light brown stripe at the level of major spine (Fig. 7e) ...... C. rafaeli n. sp. - Wings with posterior 1/2 bicolor, posterior region to first RP branch light amber, posterodistal wing region hyaline (Ardila- Camacho & García 2015, Fig. 21f); forefemur completely dark reddish brown to black ...... C. semihyalina (Le Peletier de Saint-Fargeau & Audinet-Serville, 1825) 4. Body predominantly reddish brown; pronotum with a distinct hump at midlength in lateral view (Ardila-Camacho & García 2015, Fig. 20b) ...... C. porosa Hoffman, 2002 - Body color pattern composed of a mixture of yellow and black (Ardila-Camacho & García 2015, Fig. 17); pronotum with dor- sal and ventral sides subparallel in lateral view, without a distinct hump...... C. amapaensis Penny, 1982

Genus Dicromantispa Hoffman, 2002

Type species: Mantispa sayi Banks, 1897: 23.

Dicromantispa debilis (Gerstaecker, 1888)

Mantispa debilis Gerstaecker, 1888 [“1887”]: 114. Syntypes: sex unknown, Brazil (EMAU). Mantispilla debilis var. nuda Stitz, 1913: 19. Holotype: female, Surinam (ZMB). Mantispa (Mantispilla) lineaticollis Enderlein, 1910: 348. Holotype: male, Brazil, Pará, Faro (MZPW)

New locality records. Colombia: Antioquia: Puerto Berrío, Alto de las Águilas, Hda. Manaos, 440 m, 6°27’10.09’’ N–74°36’13.74’’ W, 5–6.VII.2013, forest, light trap, Wolff-INPA (2 ♀–CEUA). Distribution. Colombia (Antioquia, Boyacá, Caldas, Córdoba, Magdalena, Meta), Brazil, Bolivia, Costa Rica, Ecuador, French Guiana, Guyana, Panama, Peru, Surinam, Trinidad, Tobago, Venezuela (Hoffman 1992, 2002; Ardila-Camacho & García 2015).

Dicromantispa gracilis (Erichson, 1839)

Mantispa gracilis Erichson, 1839: 169. Holotype: female, Brazil (ZMB). Mantispilla debilis var. nigricornis Stitz, 1913: 19. Holotype: female, Venezuela (ZMB). Mantispilla debilis var. rugicollis Stitz, 1913: 19. Holotype: female, Venezuela (ZMB).

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 313 Mantispa trilineata Navás, 1914: 230. Holotype: sex unknown, Brazil (BMNH). Mantispa bruchi Navás, 1915: 134. Holotype: female, Argentina (type depository dubious). Mantispa calceata Navás, 1917: 401. Holotype (or syntypes): male, Argentina (CN). Mantispa mista Navás, 1923: 196. Holotype: sex unknown, Argentina (MACN). Mantispa gounellei Navás, 1934: 16. Holotype: female, Brazil (MNHN).

Specimens examined. Colombia: Tolima: Icononzo, Cafrerias, 5.II.2016, attracted to light, D. Gómez (1 ♀– CAUD); Espinal, 332 m, 21.VI.1961, L. Orozco, shrubbery vegetation (1♂–CTNI); Chicoral, 400 m, 5.VII.1961, M.A. Arévalo, light trap (1♂–CTNI). Colombia: Santander: Lebrija, P.B.M., 4.VII.1968 (1♀–CTNI). New locality records. Colombia: Bolívar: Cartagena de Indias, Ciénaga de la Virgen y Juan Polo, La Boquilla, Isla del Pescador, 10°25’32’’N–75°30’31’’W, 0 m, 23.VII.2016, A. García and J. Arias (14 ♂, 5 ♀–CAUD). Distribution. Argentina, Brazil, Bolivia, Colombia (Antioquia, Bolívar, Caldas, Casanare, Chocó, Cundinamarca, Magdalena, Nariño, Santander, Tolima, Valle del Cauca), Costa Rica, Ecuador, Guyana, Panama, Uruguay, Venezuela (Hoffman 1992; Ohl 2004; Ardila-Camacho & García 2015). Comments. During a field trip to the Colombian Caribbean coast, at Ciénaga de la Virgen y Juan Polo (La Boquilla, Cartagena) A. García-García and J. Arias-Pineda carried out a collection of insects with a led light trap between 19:00 and 21:00 hours (pers. comm.). The collecting trip was made in a mangrove swamp, at 200 to 500 m from the water on July 26 2016. Shortly after the trap was set, an aggregation of around 80 specimens (males and females) of D. gracilis was observed. The specimens were seen performing erratic flights and walking on the leaves of red mangrove trees (Rhizophora mangle). Numerous clusters of stalked eggs were seen on the adaxial side of the leaves of the same plant species. Since several males and females were collected at that event, it is presumed that the aggregation was related to mating and oviposition behaviors. The specimens were collected during the rainy season.

Genus Haematomantispa Hoffman, 2002

Type species: Mantispa nubeculosa Navás, 1933: 196.

Haematomantispa is a Neotropical genus composed by two species with distribution ranging from Costa Rica to Brazil (Machado & Rafael 2010). Members of this group are easily distinguished because of their striking body coloration pattern of dark reddish (Fig. 9). The pronotum is straight and covered with scattered fine setae arising flush with the pronotal surface in lateral view. Male abdomen lacking pores; pseudopenis long and whip-like. Until now, this genus has never been recorded in Colombia, although there are reports of Haematomantispa nubeculosa Navás in Panama and H. amazonica Machado & Rafael in Brazil (Hoffman 2002; Machado & Rafael 2010). The lack of records from Colombia is undoubtedly a result of the few studies on the fauna of the group in the area. Here we recorded the genus for the first time in the country, with H. nubeculosa present in the department of Antioquia.

Haematomantispa nubeculosa (Navás, 1933) Fig. 9

Mantispa nubeculosa Navás, 1933: 196, sex not indicated. Holotype: sex unknown, Costa Rica (ZMUH).

New record. Colombia: Antioquia: Puerto Berrío, Alto de las Águilas, Hacienda Manaos. 6°27’10.09’’ N– 74°36’13.74’’W, 440 m, 5–6.VII.2013, Wolff-INPA, white light trap (1 ♀–CEUA). Diagnosis. Antenna dark brown to black except seven or eight preapical yellowish flagellomeres. Fore- and hindwing with a shaded area posterior to distal portion of CuA (Fig. 9c). Male abdomen with intersegmental membrane between tergites 4 and 5 with a membrane invaginated into a single lobed pocket, extended anteriorly 1/ 5 the length of the tergite 5. Hypomeres present as small paired circular sclerites. Pseudopenis three times as long as height of gonarcal median triangle. Female terminalia with sternite 8 posteromedially rounded. Distribution. Colombia (Antioquia), Costa Rica, Panama (Hoffman 2002). Comments. This species is found in lowlands, below 1000 m.a.s.l. from Costa Rica to northern Colombia,

314 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. probably including the Pacific region and inter-Andean valleys where tropical rain forest predominates. Temporal distribution of the adults has been recorded since January to August and November (Hoffman 2002). Immature stages and hosts are unknown.

Genus Leptomantispa Hoffman, 2002

Type species: Mantispilla pulchella Banks, 1912: 179.

The genus Leptomantispa is scarcely collected and very uncommon in museums. Species of this genus are apparently arboreal, and are likely difficult to collect. Perhaps the best way to collect specimens is by light traps. Until now the genus is composed of nine valid species occurring from southwestern Canada to northern Argentina, including the Antilles (Ohl 2004; Machado & Rafael 2010; Ardila-Camacho & García 2015; Hoffman et al. 2017). From Colombia only one species has been reported, L. hoffmani Ardila-Camacho from the department of Risaralda. The genus can be distinguished because the presence of numerous fine setae on the entire dorsal surface of the pronotum, arising flush with the pronotal surface; body color pattern consisting in mixtures of yellow and brown; male abdominal tergites 4 and 5 and sometimes 3, bearing pores; male ectoproct with ventromedial lobe not sclerotized; gonarcal membrane without scales. Female spermatheca widening distally (Hoffman 2002; Machado & Rafael 2010). Here we reported the second species from Colombia, L. axillaris (Navás), collected in the departments of Antioquia and Sucre in the norther part of the country.

Leptomantispa axillaris (Navás, 1908) Fig. 3f

Mantispa axillaris Navás, 1908: 412. Type locality: Brazil: Goiás. male and female syntypes (MNHN, MZBS).

New records. Colombia: Antioquia: Remedios, 06°54’39.8” N–74°34’06.2’’W, 520 m, 23.IV.2016, manual, A. Mejía (1 ♂–CEUA); Sucre: San Onofre, Reserva San Guaré, 0 m, 25~30.V.2009, Mercurium light trap (1 ♂– CEUA). Diagnosis. Body color pattern consisting in a mixture of yellow and brown (Fig. 3f). Foreleg with outer surface of femur light brown to dark reddish brown, inner surface generally dark brown or dark reddish brown with some yellow regions close to major spine or the row of minor spines. Wings with pterostigma reddish and subcostal space hyline (Fig. 3f). Male tergites 3–4 with two pairs of posterolateral patches of pores, each one composed of 5– 8 pores arranged as curved rows; anterolateral patches on tergites 4–5 arranged in groups of 14–26 pores, each group has one or two ovoid scars in the center. Male ectoproct with ventromedial lobe bearing 8–10 short and thick setae. Pseudopenis curved, longer than gonarcus median lobe. Pseudopenal membrane covered with minute scales. Hypomeres present as two pairs of sclerotized lateral marks on the pseudopenal membrane. Distribution. Brazil (Ohl 2004), Colombia (Antioquia, Sucre). Comments. This is a small species (forewing length, 9.6–11 mm) occurring in several states of Brazil. Here its distribution is considerably extended to the northern part of South America. Adults of this species have been collected nearly during the entire year except in January and June (Machado & Rafael 2010). Altitudinal distribution ranges from 0 to 520 m. Larval stages and hosts are completely unknown (Machado & Rafael 2010).

Genus Zeugomantispa Hoffman, 2002

Type species: Mantis minuta Fabricius, 1775: 278.

Zeugomantispa compellens (Walker, 1860)

Mantispa compellens Walker, 1860: 181. Holotype (or Syntypes), sex not indicated, Brazil, Amazonia (BMNH).

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 315 Necyla uniformis Navás, 1927: 61. Holotype (or syntypes), sex not indicated, Guatemala (CN). Mantispa parvula Penny, 1982: 458. Holotype, male, Brazil, Pará (INPA).

New locality records. Antioquia: Puerto Nare, vda. Tambores, 13.V.2013, entomological net (1 ♀–CEUA). Putumayo: Mocoa, vda. Pueblo Viejo, Fca. Villa Loca, 1°11’ N–76°38’’ W, 700 m, entomological net, 25– 26.III.2015, M. Mendoza and B. Blanco (1 ♂, 1 ♀–UNAB). Distribution. Brazil, Belize, Colombia (Antioquia, Cauca, Magdalena, Putumayo, Sucre, Valle del Cauca), Costa Rica, Guatemala, French Guyana, Honduras, Mexico, Surinam, Trinidad, Venezuela (Hoffman 1992).

Zeugomantispa minuta (Fabricius, 1775)

Mantis minuta Fabricius, 1775: 278. Holotype (or Syntypes): type locality unknown, sex unknown (BMNH). Mantis lilliputiana Olivier, 1797 [“1792”]: 640. Type locality: Surinam. Mantispa flavomaculata Latreille, 1805: 94. Type locality: Surinam. Mantis viridula Houttuyn in Stoll, 1813: 77. Type locality: Surinam. Rhaphidia margaritacea Fischer von Waldheim, 1834: 330. Holotype (or syntypes), sex unknown, Brazil (type depository unknown). Mantispa viridis Walker, 1853: 227. Holotype (or syntypes), sex not indicated, USA, Florida (BMNH). Mantispa gulosa Taylor, 1862: 494. Mantispa brevicollis Banks in Baker, 1905: 88. Mantispa pallescens Navás, 1914: 229. Holotype, male, Brazil (BMNH). Mantispilla flavescens Navás, 1914: 231. Holotype (or syntypes), male, Brazil (BMNH). Mantispilla trichostigma Navás, 1921: 51. Holotype, sex unknown, Argentina (MNHN). Mantispilla viridata Navás, 1924: 59. Holotype (or syntypes), sex unknown, Costa Rica (MNHN). Mantispilla rubricata Navás, 1924: 60. Holotype (or syntypes), sex unknown, Costa Rica (MNHN). Mantispilla flavicornis Navás, 1930: 301. Holotype (or syntypes), female, Costa Rica (ZMUH). Mantispilla taina Alayo, 1968: 13. Syntypes, sex unknown, Cuba (CZC).

New locality records. Colombia: Antioquia: Remedios, 06°54’39.8” N–74°34’06.2” W, 520 m, 23.IV.2016, manual, A. Mejía (1 ♂–CEUA); Cesar: Ecoparque Los Besotes, 10°34’36” N–73°16’33” W, 15.II.2009, A. Morales, captura manual (1 ♂–CEUA). Distribution. Argentina, Bahamas, Belize, Colombia (Antioquia, Bolívar, Casanare, Cundinamarca, Magdalena, Meta), Costa Rica, Cuba, Ecuador, El Salvador, Guatemala, Honduras, Hispaniola, Mexico, Nicaragua, Panama, Peru, Suriname, Uruguay, USA, and Venezuela (Hoffman 1992).

Zeugomantispa virescens (Rambur, 1842)

Mantispa viridula Erichson, 1839: 170. Syntypes, sex unknown, Brazil (ZMB, MCZ) Mantispa virescens Rambur, 1842: 433. Holotype, male, type locality unknown (ISNB). Mantispilla punctata Stitz, 1913: 20. Syntypes, female, Brazil (ZMB). Mantispilla punctata var. major Stitz, 1913: 20. Holotype, sex unknown, Brazil (ZMB). Mantispilla stigmata Stitz, 1913: 20. Holotype, male, Brazil (ZMB). Mantispa viridis Stitz, 1913: 29. Syntypes, females, Paraguay (ZMB).

Specimens examined. Colombia: Antioquia: Bello, 1.X.1958, C. Ríos, on Zea maiz (1 ♂–CTNI); same data but 29.XII.1953, C. Carmona, on Crotalaria (1 ♂–CTNI); Ciudad Bolívar, vra. La Linda, fca. Pamplonita, 14.I.2004, white light trap, H. Vargas (1 ♀–CEUA); Puerto Berrío, Alto de las Águilas, Hda. Manaos, 400–500 m, 6°27'10.09''N–74°36'13.74''W, 06.VII.2013, forest, GEUA (4 ♂, 1 ♀–CEUA); Sucre: San Onofre, Rva. Sanguaré, 6º27’15.5” N–74º51’28.7” W, 22–23.III.2004, grassland, mercury light trap, GEUA (1 ♂–CEUA); San Onofre, Rva. Sanguaré, 6º27’15.5” N–74º51’28.7” W, 25–30.V.2009, forest, mercury light trap, GEUA (8 ♂, 2 ♀–CEUA); San Onofre, Rva. Sanguaré, P. La Cofrería, 11–15.II.2010, mercury light trap, GEUA (1 ♂–CEUA); San Onofre, Rva. Sanguaré, 6º27’15.5” N–74º51’28.7” W, 21.VII.2009, forest, mercury light trap, J.D. Sáchez; A. Montoya (2 ♂, 3 ♀–CEUA); Valle del Cauca: Tablones, 7.I.1959, J.F.G. Clarke (1 ♀–CTNI).

316 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. FIGURE 7. Climaciella rafaeli n. sp.: a, fore- and hindwing; b, head and thorax, lateral view; c, head, frontal view; d, head and thorax, dorsal view; e, forefemur, outer surface; f, same, ventrolateral view.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 317 FIGURE 8. Climaciella rafaeli n. sp.: a, male genitalia, dorsolateral view; b, same, ventral view; c, pregenital abdominal apparatus, dorsal; d, e, male genitalia, dorsal view; f, male genitalia, ventral view; g, sternite 9, ventral view; h, gonarcus, ventral view. Abbreviations: g, gonarcus; gcx9, gonocoxite 9; hpm, hipomere; med, mediuncus; po, pores; ppm, pseudopenal membrane; psp, pseudopenis; sc, scar; t3‒t6, third to sixth abdominal tergites; vml, ventromedial lobe. Scale bars: 0.1 mm, except for c.

318 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. Distribution. Argentina, Brazil, Colombia (Antioquia, Bolívar, Caldas, Cundinamarca, Huila, Magdalena, Meta, Quindío, Risaralda, Sucre, Tolima, Valle del Cauca), Costa Rica, Ecuador, Guyana, French Guiana, Mexico, Nicaragua, Panama, Peru, Surinam, Tobago, Trinidad, Uruguay, Venezuela (Hoffman 1992).

FIGURE 9. Haematomantispa nubeculosa: a, female habitus; b, forefemur, outer (left) and inner (right) surfaces; c, pterothorax and abdomen, lateral view; d, head, frontal view.

Updated key to genera of Colombian Mantispinae (After Ardila-Camacho & García 2015, modified to include the newly recorded genus Haematomantispa)

1. Antennal flagellomeres wider than long in frontal view (Fig. 7c); wings with anterior 1/2 light to dark amber (Fig. 7a); hind- wing with crossvein between CuA and A1 (Fig. 7a) ...... Climaciella - Antennal flagellomeres as wide as long in frontal view (Fig. 4c); wings usually completely hyaline (Fig. 4a); hindwing without crossvein between CuA and A1 (Fig. 4a) ...... 2 2. Pronotum with numerous prominent setae over entire length in lateral view (Hoffman 2002, Figs. 608, 609)...... 3 - Pronotum with scattered fine setae in lateral view (Ardila-Camacho & García 2015, Fig. 22a; Hoffman 2002, Figs. 606, 607) ...... 5 3. Body predominantly light yellow or green (Machado & Rafael 2010, Figs. 27, 28); pronotum with numerous setae arising from protuberant bases (Machado & Rafael 2010, Fig. 29c)...... Zeugomantispa - Body with different color pattern; pronotum with setae arising flush with pronotal surface (Hoffman 2002, Fig. 608) ...... 4 4. Body mostly dark red (Fig. 9a, c); forewing with an amber infuscation posterior to distal portion of CuA (Machado & Rafael 2010, Fig. 17f); male abdominal tergites without pores ...... Haematomantispa - Body with a mixture of brown and yellow or tan (Fig. 6; Ardila-Camacho & García 2015, Fig. 23); forewing without an amber infuscation posterior to distal portion of CuA; male abdominal tergites with pores ...... Leptomantispa 5. Head with inverted V-shaped mark directly beneath antennal insertion (Ardila-Camacho & García 2015, Fig. 1c); body color- ation mottled, camouflage-like, consisting of green or light yellow and brown (Fig. 3d); hypomeres long (Machado & Rafael 2010, Fig. 4e) ...... Buyda - Head with longitudinal or transverse stripes beneath antennae; body with varied color pattern; hypomeres short ...... 6 6. Body mostly brown or tan (Machado & Rafael 2010, Fig. 5); mesonotum with paired longitudinal yellow stripes laterally; forewing sometimes amber at base (Machado & Rafael 2010, Fig. 13f); male ectoproct with ventromedial lobe completely sclerotized ...... Dicromantispa - Body generally yellow and black (Ardila-Camacho & García 2015, Fig. 22); mesonotum with paired yellow stripes angled

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 319 from wings bases to scutellum (Ardila-Camacho & García 2015, Fig. 22e); forewing with basal third of rarp1 (first anterior radial cell) amber (Ardila-Camacho & García 2015, Fig. 22f); male ectoproct with ventromedial lobe partially sclerotized ...... Entanoneura

Discussion

In the review of the Mantispidae of Colombia, 24 named species were recognized to occur in the country. Within these reported taxa, four were described as new to science; two were classified as incertae sedis and one as an erroneous record. Thus, the family was represented by a total of 21 nominal species: one in Drepanicinae (a record that needs to be corroborated, because the collecting data were questionable); 13 species in Mantispinae, and seven species in Symphrasinae (Ardila-Camacho & García 2015). With the data reported here, the number of species of Colombia increases to 26 named species (Table 2). Within the new additions, four species belong to Mantispinae, Leptomantispa axillaris (widely distributed in Brazil), Haematomantispa nubeculosa, that represents the first record of the genus in Colombia (previously known from Central America), and the new species of Buyda immaculata and Climaciella rafaeli; the former species was previously described by Hoffman (1992), but was never formally published. The fifth species corresponds to the subfamily Symphrasinae, the new Amazonian species Trichoscelia umbrata. Although most of the Colombian Museums were revised in the work of Ardila- Camacho & García (2015), specimens from other museums were not available for the study, so the addition of new taxa to the Colombian diversity of Mantispidae was expected. Based on the recent revision of the specimens deposited in CEUA, several species are considered here as new records from Colombia. Many of the reported species are mainly distributed in the department of Antioquia, region where most of the material has been collected. Other specimens deposited in the collections of UNAB, CAUD, and ANDES-E, were also included. Among the new distributional records of Colombian Mantispidae, Dicromantispa debilis known from Boyacá, Caldas, Magdalena and Meta, is here reported from Antioquia, and Dicromantispa gracilis, one of the most common mantispid species in Colombia (Antioquia, Bolívar, Caldas, Casanare, Magdalena, Nariño, Santander, Tolima, and Valle del Cauca), is newly reported from Bolivar. Zeugomantispa minuta previously known from Bolívar, Casanare, Cundinamarca, and Magdalena, is here reported from Antioquia and Cesar, while Z. compellens known from Cauca, Magdalena, Sucre, and Valle del Cauca is reported to occur in Antioquia and Putumayo. Within Symphrasinae, Plega hagenella accounts with records from Antioquia, Chocó, Cundinamarca, Magdalena, Meta, and Santander and is here reported from Caquetá. From the 32 departments conforming the Colombian territory, the best represented by Mantispidae corresponds to Antioquia, where 13 species are known to occur. Magdalena reports nine species and Cundinamarca, Meta, and Valle del Cauca have six species each, while Bolivar is represented by four species. The remainder departments are only represented by one to three species, except for Atlantico, Guainia, Guaviare, La Guajira, Norte de Santander, San Andrés y Providencia, and Vaupés, where the records of Mantispidae are lacking (Table 2). This is certainly an artifact due to absence of studies focusing on this insect group, and in general, there is a remarkable contrast with other countries, such as Brazil, where the group has been well studied. For instance, in a single locality of Manaus (Amazonas), 143 specimens belonging to 12 species were reported during a year of collection (Machado 2007). In contrast, in the Colombian department of Amazonas only one species has been reported (Ardila-Camacho & García 2015). Thereby, a vast area of some of the most biodiverse regions of the world such as the Choco, Amazonian rainforest, or the mountain ranges of the Andes and their valleys are still poorly represented by this family, so new records and new species are expected to be discovered as systematic collections are performed. The relatively low number of records of Mantispidae are generally erroneously attributed to the rareness of the group. However, several studies have demonstrated that mantispids are certainly a relatively common group, with ecological parameters such as abundance and a very variable diversity, depending on the biogeographical region. For Mantispinae, Machado (2007) demonstrated that the abundance of the Amazonia species seems to be inverse to the species richness, while New & Haddow (1973) showed a higher abundance but a low diversity in a similar study in the Afrotropical region (e.g., 1106 specimens belonging to six species during a year of collection). In both cases, the number of captured specimens was considerably high. Also, some studies carried out on Nearctic species have showed a high rate of parasitism from mantispids on different spider families (Rice & Peck 1991; Redborg & Redborg 2000). In studies focused on Symphrasinae something similar happens, where numerous collections of the

320 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. TABLE 2. Updated check-list of Colombian Mantispidae (modified from Ardila-Camacho & García 2015), new records are in bold letters. Subfamily Genus/species Geographical records in Colombia (Departments) Mantispinae Buyda Navás, 1926 Buyda immaculata n. sp. Antioquia, Sucre Buyda phthisica (Gerstaecker, 1885) Antioquia, Cundinamarca, Meta, Valle del Cauca Climaciella Enderlein, 1910 Climaciella amapaensis Penny, 1992 Risaralda Climaciella obtusa Hoffman, 2002 Magdalena, Valle del Cauca Climaciella porosa Hoffman, 2002 Valle del Cauca Climaciella rafaeli n. sp. Antioquia Climaciella semihyalina (Le Peletier de Saint- Putumayo Fergeau y Audinet-Serville, 1825) Dicromantispa Hoffman, 2002 Dicromantispa debilis (Gerstaecker, 1888) Antioquia, Boyacá, Caldas, Córdoba, Magdalena, Meta Dicromantispa gracilis (Erichson, 1839) Antioquia, Bolívar, Caldas, Casanare, Magdalena, Nariño, Santander, Tolima, Valle del Cauca Dicromantispa moulti (Navás, 1909) Amazonas Entanoneura Enderlein, 1910 Entanoneura batesella (Weswood, 1867) Arauca, Magdalena Haematomantispa Hoffman, 2002 Haematomantispa nubeculosa (Navás, 1933) Antioquia Leptomantispa Hoffman, 2002 Leptomantispa axillaris (Navás, 1908) Antioquia, Sucre Leptomantispa hoffmani Ardila-Camacho, 2015 Risaralda Zeugomantispa Hoffman, 2002 Zeugomantispa compellens (Walker, 1860) Antioquia, Cauca (Gorgona island), Magdalena, Sucre, Putumayo, Valle del Cauca Zeugomantispa minuta (Fabricius, 1775) Antioquia, Bolívar, Casanare, Cesar, Cundinamarca, Magdalena Zeugomantispa virescens (Rambur, 1842) Antioquia, Bolívar, Caldas, Cundinamarca, Huila, Magdalena, Meta, Quindío, Risaralda, Sucre, Tolima, Valle del Cauca Symphrasinae Anchieta Navás, 1909 Anchieta eurydella (Westwood, 1867) Meta Anchieta fasciatella (Westwood, 1867) Bolívar, Magdalena, Santander Anchieta remipes (Gerstaecker, 1888) Cundinamarca Plega Navás, 1927 Plega hagenella (Westwood, 1867) Antioquia, Caquetá, Chocó, Cundinamarca, Magdalena, Meta, Santander Trichoscelia Westwood, 1852 Trichoscelia umbrata n. sp. Caquetá Trichoscelia andina Ardila-Camacho, 2015 Cundinamarca Trichoscelia gorgonensis Ardila-Camacho, 2015 Cauca (Gorgona island) Trichoscelia karijona Ardila-Camacho, 2015 Caquetá, Meta, Putumayo, Vichada Drepanicinae Gerstaeckerella Enderlein, 1910 Gerstaeckerella gigantea Enderlein, 1910 Antioquia*

*Record need to be confirmed.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 321 North American species Plega dactylota Rehn, aggregations of the Neotropical Trichoscelia santareni (Navás) and massive parasitism of Plega hagenella on stingless bees have been documented (Rice 1987; Dejean & Canard 1990; Maia-Silva et al. 2013). It would be very useful to perform systematic collecting using different light- and flight intercepting traps in the different Colombian biogeographic provinces, in order to have a better estimate of the species richness of Mantispidae in the country.

Acknowledgements

We offer our sincere thanks to the curators of the Museums revised, Erika Valentina Vergara (Facultad de Agronomía, Universidad Nacional de Colombia, Bogotá) and Colección Entomológica “Luis María Murillo”, Corpoica, Mosquera, Cundinamarca), Francisco Serna (Facultad de Agronomía, Universidad Nacional de Colombia, Bogotá), Alexander García-García (Universidad Distrital de Bogotá), and Emilio Realpe (Universidad de los Andes). Sincere thanks to John Albeiro Quiroz and Guillermo Guarín Candamil (Museo Entomlógico “Francisco Luis Gallego”, Universidad Nacional de Colombia, sede Medellín) for their help and disinterested kindness during the visit of the first author to their institution. We also thank Isabel Carmona Gallego who critically reviewed the English. The first author wants to thank to Neusa Hamada, Jean Marcelle Carvalcante and Thiago Mahlmann (Instituto Nacional de Pesquisas da Amazônia-INPA) for their kind help, information on the type material deposited in INPA, and for sending photographs of the type specimen of Trichoscelia anae. Thanks to Kevin M. Hoffman (California Department of Food and Agriculture, Sacramento) for his valuable comments and support during the evelopment of this work. Sincere thanks to David Bowles (Missouri State University, USA) for his steady support and for providing photographs and information on Neotropical Symphrasinae. Thanks to Amoret Spooner (Oxford University Museum, UK) for kindly provide photographs of the type material of Trichoscelia iridella. AAC offers his gratitude to the Departamento de Ciencias Biológicas, Universidad de los Andes, for the financial support.

References

Ardila-Camacho, A. & García, A. (2015) Mantidflies of Colombia (Neuroptera, Mantispidae). Zootaxa, 3937 (3), 401–455. https://doi.org/10.11646/zootaxa.3937.3.1 Austin, A.D. (1985) The function of spider egg sacs in relation to parasitoids and predators, with special reference to the Australian fauna. Journal of Natural History, 19, 359‒376. https://doi.org/10.1080/00222938500770261 Aspöck, U., Plant, J.D. & Nemeschkal, H.L. (2001) Cladistic analysis of Neuroptera and their systematic position within Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology, 26, 73‒86. https://doi.org/10.1046/j.1365-3113.2001.00136.x Aspöck, U. & Aspöck, H. (2007) Verbliebene Vielfalt vergangener Blüte. Zur Evolution, Phylogenie und Biodiversität der Neuropterida (Insecta: Endopterygota). Denisia, 20, 451–516. Aspöck, U. & Aspöck, H. (2008) Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Systematic Entomology, 33, 97–127. https://doi.org/10.1111/j.1365-3113.2007.00396.x Aspöck, U. & Mansell, M.W. (1994) A revision of the family Rhachiberothidae Tjeder, 1959, stat. n. (Neuroptera). Systematic Entomology, 19, 181–206. https://doi.org/10.1111/j.1365-3113.1994.tb00587.x Breitkreuz, L.C.V., Winterton, S.L. & Engel, M.S. (2017) Wing Tracheation in Chrysopidae and Other Neuropterida (Insecta): A Resolution of the Confusion about Vein Fusion. American Museum Novitates, 3890, 1‒44. https://doi.org/10.1206/3890.1 Buys, S.C. (2008) Observations on the biology of Anchieta fumosella (Westwood 1867) (Neuroptera: Mantispidae) from Brazil. Tropical Zoology, 21, 91‒95. Dejean, A. & Canard, M. (1990) Reproductive behaviour of Trichoscelia santareni (Navás) (Neuroptera: Mantispidae) and parasitization of the colonies of Polybia diguetana R. du Buysson (Hymenoptera: Vespidae). Neuroptera International, 6, 19‒26. Dorey, J.B. & Merrit, D.J. (2017) First observations on the life cycle and mas eclosion events in a mantis fly (Family Mantispidae) in the subfamily Drepanicinae. Biodiversity Data Journal, 5 (e21206), 2‒12. Frantsevich, L. (1998) The Coxal Articulation of the Insect Striking Leg: A Comparative Study. Journal of Morphology, 236,

322 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL. 127‒138. https://doi.org/10.1002/(SICI)1097-4687(199805)236:2%3C127::AID-JMOR4%3E3.0.CO;2-2 Hoffman, K.M. (1992) Systematics of the Mantispinae (Neuroptera:Mantispidae) of North, Central, and South America. Unpublished Ph. D. dissertation, Clemson University, Clemson, South Carolina, 501 pp. Hoffman, K.M. (2002) Family Mantispidae. In Penny, N. D. (ed.), A guide to the lacewings (Neuroptera) of Costa Rica. Proceedings of the California Academy of Sciences, 53 (12), 251–275. Hoffman, K.M. & Brushwein, J.R. (1992) Descriptions of the larvae and pupae of some North American Mantispinae (Neuroptera: Mantispidae) and development of a system of larval chaetotaxy for Neuroptera. Transactions of the American Entomological Society, 118, 159‒196. Hoffman, K.M., Flint, O.S. Jr. & Perez-Gelabert, D.E. (2017) The Mantispidae of the West Indies with special reference to the Dominican Republic (Neuroptera: Mantispidae). Insecta Mundi, 0559, 1‒15. Hook, A.W., Oswald, J.D. & Neff, J.L. (2010) Plega hagenella (Neuroptera: Mantispidae) parasitism of Hylaeus (Hylaeopsis) sp. (Hymenoptera: Colletidae) reusing nests of Trypoxylon manni (Hymenoptera: Crabronidae) in Trinidad. Journal of Hymenoptera Research, 19, 77‒83. Kral, K. (2013) Vision in the mantispid: a sit-and-wait and stalking predatory insect. Physiological Entomology, 38, 1–12. https://doi.org/10.1111/phen.12004 Lambkin, K.J. (1986) A revision of the Australian Mantispidae (Insecta: Neuroptera) with a contribution to the classification of the family I. General and Drepanicinae. Australian Journal of Zoology, Supplement Series, 116, 1–142. https://doi.org/10.1071/AJZS116 Liu, X.Y., Winterton, S.L., Chao, W., Ross, P. & Ohl, M. (2015) A new genus of mantidflies discovered in the Oriental region, with a higher-level phylogeny of Mantispidae (Neuroptera) using DNA sequences and morphology. Systematic Entomology, 40 (1), 183–206. https://doi.org/10.1111/syen.12096 Machado, R.J.P. (2007) Taxonomia e sazonialidade das espécies de Mantispidae (Insecta: Neuroptera) da Amazônia Brasileira. Master's thesis. Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, xvi + 127 pp. Machado, R.J.P. & Rafael, J.A. (2010) Taxonomy of the Brazilian species previously placed in Mantispa Illiger, 1798 (Neuroptera: Mantispidae), with the description of three new species. Zootaxa, 2454, 3–59. MacLeod, E.G. & Redborg, K.E. (1982) Larval Platymantispinae Mantispids (Neuroptera: Planipennia): possibly a subfamily of generalist predators. Neuroptera international, 2 (1), 37–41. Maia-Silva, C., Hrncr, M., Koedam, D., Machado, R.J.P. & Imperatriz-Fonseca, V.L. (2013) Out with the garbage: the parasitic strategy of the mantisfly Plega hagenella mass-infesting colonies of the eusocial bee Melipona subnitida in northeastern Brazil. Naturwissenschaften, 100, 101–105. https://doi.org/10.1007/s00114-012-0994-1 Monserrat, V.J. (1985) Lista de los tipos de Mecoptera y Neuroptera (Insecta) de la colección L. Navás, depositados en el Museo de Zoología de Barcelona. Miscellánia Zoológica, 9, 233‒243. Monserrat, V.J. (2014) Los mantíspidos de la Península Ibérica y Baleares (Insecta, Neuropterida, Neuroptera, Mantispidae). Graellsia, 70 (2), 1‒52. https://doi.org/10.3989/graellsia.2014.v70.115 Navás, L. (1914) Névroptères nouveaux de l'Amerique du Nord. IIème série. Entomologische Zeitschrift, Frankfurt am Main, 28, 18‒20, 25‒26. Navás, L. (1926) Insectos exóticos Neurópteros y afines. Brotéria (Zoológica), 23,79–93. Navás, L. (1933) Insectos suramericanos. Septima [VII] serie. Revista de la Real Academia de Ciencias Exactas Fisicas y Naturales de Madrid, 30, 303‒314. New, T.R. (1989) Planipennia, Lacewings. Handbuch der Zoologie, Vol. 4 (Arthropoda: Insecta). Part 30. Walter de Gruyter, Berlin, 132 pp. New, T.R. & Haddow, A.J. (1973) Nocturnal flight activity of some African Mantispidae (Neuroptera). Journal of Entomology, London, (A), 47,161–168. https://doi.org/10.1111/j.1365-3032.1973.tb00021.x Ohl, M. (2004) Annotated catalog of the Mantispidae of the world (Neuroptera). Contributions on Entomology, International, 5 (3), 131–262. Ohl, M. (2005) Toward a global inventory of Mantispidae—the state-of-the-art in mantispid taxonomy. In: Pantaleoni, R.A., Letardi, A. & Corazza, C. (Eds.), Proceedings of the Ninth International Symposium on Neuropterology (20–23 June 2005, Ferrara, Italy). Annali del Museo Civico di Storia Naturale di Ferrara, 8, pp. 79–86. Ohl, M. (2011) Aboard a spider—a complex developmental strategy fossilized in amber. Naturwissenschaften, 98, 453–456. https://doi.org/10.1007/s00114-011-0783-2 Opler, P.A. (1981) Polymorphic Mimicry of Polistine Wasps by a Neotropical Neuropteran. Biotropica, 13, 165−176. https://doi.org/10.2307/2388121 Parker, F.D. & Stange, L.A. (1965) Systematic and biological notes on the tribe Platymantispini (Neuroptera: Mantispidae) and the description of a new species of Plega from Mexico. Canadian Entomologist, 97, 604‒612. https://doi.org/10.4039/Ent97604-6 Penny, N.D. (1982) Neuroptera of the Amazon basin. Part 6. Mantispidae (1). Acta Amazônica, 12 (2), 415–463.

NEW SPECIES AND NEW RECORDS OF NEOTROPICAL MANTIDFLIES Zootaxa 4413 (2) © 2018 Magnolia Press · 323 https://doi.org/10.1590/1809-43921982122415 Penny, N.D. & da Costa, C.A. (1983) Mantispídeos do Brasil (Neuroptera: Mantispidae). Acta Amazônica, 13 (3–4), 601–687. https://doi.org/10.1590/1809-439219831334601 Redborg, K.E. (1998) Biology of the Mantispidae. Annual Review of Entomology, 43, 175−194. https://doi.org/10.1146/annurev.ento.43.1.175 Redborg, K.E. & Redborg, A.H. (2000) Resource partitioning of spider hosts (Arachnida, Araneae) by two mantispid species (Neuroptera, Mantispidae) in an Illinois woodland. Journal of Arachnology, 28, 70–78. https://doi.org/10.1636/0161-8202(2000)028[0070:RPOSHA]2.0.CO;2 Reynoso-Velasco, D. & Contreras-Ramos, A. (2008) Mantispidae (Neuroptera) of Mexico: Distribution and key to genera. Annals of the Entomological Society of America, 101 (4), 703–712. https://doi.org/10.1093/aesa/101.4.703 Rice, M.E. (1987) Morphological variation in Plega dactylota (Neuroptera: Mantispidae) and status of its subspecies. Journal of the Kansas Entomological Society, 60, 341−344. Rice, M.E. & Peck, W.B. (1991) Mantispa sayi (Neuroptera: Mantispidae) parasitism on spiders (Araneae) in Texas, with observations on oviposition and larval survivorship. Annals of the Entomological Society of America, 84, 52−57. https://doi.org/10.1093/aesa/84.1.52 Snyman, L.P., Ohl, M., Mansell, M.W. & Scholtz, C.H. (2012) A revision and key to the genera of Afrotropical Mantispidae (Neuropterida, Neuroptera), with the description of a new genus. ZooKeys, 184, 67−93. https://doi.org/10.3897/zookeys.184.2489 Tauber, C.A., Legrand, J., Albuquerque, G.S., Ohl, M., Tauber, A. & Tauber, M.J. (2017) Navás’ specimens of Mantispidae (Neuroptera) in the Museum National D’Histoire Naturelle, Paris. Proceedigs of the Entomological Society of Washington, 119 (2), 239‒263. https://doi.org/10.4289/0013-8797.119.2.239 Willmann, R. (1990) The phylogenetic position of the Rhachiberothinae and the basal sister-group relationships within the Mantispidae (Neuroptera). Systematic Entomology, 15, 253–265. https://doi.org/10.1111/j.1365-3113.1990.tb00316.x Winterton, S.L., Lemmon, A.R., Gillung, J.P., Garzon, I.J. Badano, D., Bakkes, D.K., Breitkreuz, L.C.V., Engel, M.S., Lemmon, E.M., Liu, X., Machado, R.J.P., Skevington, J.H. & Oswald, J.D. (2017) Evolution of lacewings and allied orders using anchored phylogenomics (Neuroptera, Megaloptera, Raphidioptera). Systematic Entomology. [published online] https://doi.org/10.1111/syen.12278

324 · Zootaxa 4413 (2) © 2018 Magnolia Press ARDILA-CAMACHO ET AL.