Insecta: Neuropterida) of Colombia

AQUATIC INSECTS 2018, VOL. 39, NO. 2-3, 297–353 https://doi.org/10.1080/01650424.2018.1500001 The Megaloptera (Insecta: Neuropterida) of Colombia aÃ b Adrian Ardila-Camacho and Atilano Contreras-Ramos aEscuela de Posgrado en Ciencias Biologicas, Departamento de Ciencias Biologicas, Universidad de los Andes, Bogota, Colombia; bDepartamento de Zoologıa, Instituto de Biologıa, Universidad Nacional Autonoma de Mexico, Ciudad Universitaria, Ciudad de Mexico, Mexico ABSTRACT ARTICLE HISTORY The knowledge of Megaloptera (Neuropterida) fauna of Colombia Received 11 January 2018 is updated. Based on the specimens studied and literature records, Accepted 2 April 2018 it was determined that 18 species grouped into three genera First published online 14 December 2018 (Ilyobius Enderlein, 1910, Chloronia Banks, 1908, and Corydalus Latreille, 1802) and two families (Sialidae and Corydalidae) are KEYWORDS recognized to occur in the country. Within the provided records, Dobsonflies; alderflies; two new species of Corydalus, Corydalus liui sp. n. and Corydalus Corydalus; new species; sophiae sp. n., are described and illustrated. Last larval instar of faunistics; Colombia C. armatus Hagen, 1861 and C. liui sp. n. are described and illustrated. Comments about the biology and distribution of the majority of the species are included. Illustrations of the external morphology and genital structures of selected species are also provided. Keys for identification of adults and larvae accounting with descriptions of Colombian Megaloptera are presented. Introduction The order Megaloptera is an ancient lineage of insects with phylogenetic origins near the base of the Holometabola, that together with the Neuroptera and Raphidioptera constitute the superorder Neuropterida (Aspock,€ Plant, and Nemeschkal 2001). This superorder is considered as a relict comprising approximately 6500 species worldwide, from which Megaloptera has approximately 380 representatives grouped in the families Sialidae and Corydalidae (Winterton, Hardy, and Wiegmann 2010; Liu, LU,€ Aspock,€ Yang, and Aspock€ 2015a). Estimates of phylogenetic relationships of Megaloptera have been controversial both within the order and in the context of Neuropterida (Winterton et al. 2010; Aspock,€ Haring, and Aspock€ 2012). Traditional hypotheses proposed Megaloptera as sister to Raphidioptera (Kristensen 1999; Whiting 2001; Beutel et al. 2011), but recently molecular and morphological evidence has accumulated that supports the position of Raphidioptera as sister of Neuroptera þ Megaloptera (Aspock€ et al. 2001; Haring and Aspock€ 2004; Aspock€ and Aspock€ 2008; Beutel and Friedrich 2008; Wang, Liu, Winterton, and Yang 2012; CONTACT Adrian Ardila-Camacho [email protected] Escuela de Posgrado en Ciencias Biologicas, Departamento de Ciencias Biologicas, Universidad de los Andes, Bogota, Colombia Ã Present address: Doctorado en Ciencias Biologicas, sede Instituto de Biologıa-UNAM, Ciudad Universitaria, Ciudad de Mexico, Mexico ß 2018 Informa UK Limited, trading as Taylor & Francis Group Published online 20 Dec 2018 298 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS Zhao, Liu, and Yang 2014; Wang et al. 2017; Winterton et al. 2018). Furthermore, a monophyletic Megaloptera with Sialidae as sister of Corydalinae þ Chauliodinae, is a well-supported hypothesis under different approaches (Aspock€ et al. 2001; Aspock€ and Aspock€ 2008; Wang et al. 2012; Zhao et al. 2014; Liu et al. 2015a; Wang et al. 2017; Winterton et al. 2018). The Megaloptera are distributed in all biogeographical realms, with isolated and often discontinuous distribution patterns, which are product of a long evolutionary history (Cover and Resh 2008). Oldest fossils of the group are known from the Permian, and the divergence between Sialidae and Corydalidae was estimated to have occurred at the beginning of the Jurassic (Jepson and Heads 2016; Winterton et al. 2018). With a single extant subfamily (Sialidinae), the family Sialidae is distributed worldwide, and is represented by 75 species (Liu, Hayashi, and Yang 2015b). Corydalidae is divided into two subfamilies, Corydalinae and Chauliodinae (Liu et al. 2015a). The Chauliodinae, or fishflies, have a disjunct distribution in Asia, Australia, North America, southern South America, South Africa, and Madagascar, with approximately 128 described species (Liu, Wang, Shih, Ren, and Yang 2012). Corydalinae, or dobsonflies, contains about 160 species, mainly distributed in the southern hemisphere, although there are a few representatives in North America and Asia (Cover and Resh 2008; Jiang, Yang, Yang, and Liu 2016). The Neotropical Region is represented by 10 species of Sialidae, 10 species and one subspecies of Chauliodinae, and 56 species of Corydalinae (Contreras-Ramos 2005; Cover and Resh 2008; Cardoso-Costa, Azevêdo, and Ferreira 2013; Liu et al. 2015b). The Neotropical fauna of Megaloptera is well documented, and taxonomic revi- sions with illustrations, keys, and descriptions based mainly on male features are available (Penny and Flint 1982; Glorioso and Flint 1984; Contreras-Ramos 1998). Females are often identified based on mandible morphology, colour pattern, and distribution but female genitalia are quite uniform between species, and so species iden- tifications may be unreliable. Immature stages have been scarcely studied, and thereby the vast majority of species need to be identified by rearing mature larvae, or by using methods such as DNA barcoding (Contreras-Ramos 1999a; Jung, Vshivkova, and Bae 2015). Available keys and/or descriptions of larval Megaloptera of the Neotropics are restricted to the fauna of a few regions, such as Chile, Mexico, and the Brazilian Amazonia (Evans 1972; Flint 1973; Penny and Flint 1982; Contreras- Ramos and Harris 1998; Azevêdo 2003, 2009; Azevêdo and Hamada 2006, 2007, 2014). Thus, in most of the Neotropics, larval Megaloptera are largely uncharacter- ized, despite the high species diversity found in this region (Cover and Resh 2008) and the high abundance of larvae in many streams and rivers. Prior to this article, the Colombian fauna of Megaloptera was represented by 11 species of Corydalus Latreille, 1802, two of Chloronia Banks, 1908 and two of Ilyobius Enderlein, 1910 (Contreras-Ramos 1998, 2005; Ardila-Camacho 2014; Liu, Hayashi, and Yang 2015c). There have been no studies on immature stages of Colombian Megaloptera, but the genera and a few species shared with Brazil may be identified following Azevêdo and Hamada (2014). In view of the importance of Megaloptera to aquatic ecosystems and for the assessment of water quality, the purpose of this study is to update the taxonomic knowledge of the group in Colombia, and to provide a AQUATIC INSECTS 299 check-list and taxonomic keys to the larvae and adults of the genera and species known to occur there. Also, two new species of Corydalus, and the last instar larva of Corydalus liui sp. n., and Corydalus armatus Hagen, 1861 are described and illustrated. Diagnosis and distribution for each species are also given. Material and methods Immature stages of Megaloptera were collected by hand or using a D-frame net. The collecting sites are located on the western slope of the Eastern Cordillera of Colombia. Last instar larvae of two Corydalus species were captured during December 2014, February, April to June, October, and December 2015, and January 2016 in a small stream in the vicinity of the Chipata Municipality, Santander Department (60304500N, 733802400W; above 1700 m above see level (a.s.l.)). In the field, the larvae were placed in zippered plastic bags with small holes containing small stones and moist mosses; subsequently the bags were placed inside of a plastic box without water and allowing aeration. In the laboratory, the larvae were reared indi- vidually inside of glass bottles submerged in a large aquarium (60 Â 24 Â 40 cm) with constant and abundant air flux provided by an air pump of four outputs. Each bottle was set with sand and small stones and was closed with a plastic net. The aquarium was maintained at 23 C and 12:00/12:00 hours (light/darkness). Larvae were fed every three days with house crickets (Acheta domesticus (Linnaeus 1758)), earthworms (Eisenia fetida (Savigny 1826)), small fishes (Poecilia sp.), and brine shrimp (Artemia salina (Linnaeus 1758)). Body measurements were taken with a digital caliper (0.01 mm accuracy). Mature larvae were induced to pupate by placing them into individual con- tainers filled with substrate taken from the collecting sites into which a cavity was made and then covered by a flat stone (Contreras-Ramos 1999a). The substrate was regularly moistened. All the specimens induced to pupation were maintained at 26 C. Selected late instar larvae were anesthetized at À5 C for 30 minutes, then were fixed with a syringe injection through the mouth of a solution of 10% formalin in phosphate buffer (pH 7.4). After 12 hours in the fixing agent, the larvae were washed with phosphate buffer solution at pH 7.4, and then preserved in 70% ethyl alcohol. Morphological descriptions were made using a Zeiss Stemi 2000 stereomicroscope, adapted with an AxioCam ERc 5s digital camera. Adults obtained in the laboratory (from successful last instar pupations) were killed in 80% ethanol one day after the emergence, following hardening of the cuticle. Then, the specimens were pinned and their wings extended. Some individuals were preserved and stored in 96% ethanol. Species identification was made by examination of the genitalia. The last four abdom- inal segments

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