AQUATIC INSECTS 2018, VOL. 39, NO. 2-3, 297–353 https://doi.org/10.1080/01650424.2018.1500001
The Megaloptera (Insecta: Neuropterida) of Colombia
a b Adrian Ardila-Camacho and Atilano Contreras-Ramos aEscuela de Posgrado en Ciencias Biologicas, Departamento de Ciencias Biologicas, Universidad de los Andes, Bogot a, Colombia; bDepartamento de Zoolog ıa, Instituto de Biolog ıa, Universidad Nacional Autonoma de M exico, Ciudad Universitaria, Ciudad de M exico, 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 fami- lies 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, Bogot a, Colombia Present address: Doctorado en Ciencias Biologicas, sede Instituto de Biolog ıa-UNAM, Ciudad Universitaria, Ciudad de M exico, 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, Azev^edo, 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 dis- tribution 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; Azev^edo 2003, 2009; Azev^edo 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 Azev^edo 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 illus- trated. 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 Chipat a Municipality, Santander Department (6 0304500N, 73 3802400W; 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 24 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 5 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 of the males were dissected. These were then cleared in 10% potassium hydroxide (KOH) overnight. After treatment with KOH, the genitalia were washed with distilled water, 10% acetic acid, and 70% ethanol. These structures were pre- served in glass tubes filled with glycerine and pinned below the respective specimen. Morphological terminology of the larvae follows that of New and Theischinger (1993), Contreras-Ramos and Harris (1998), and Azev^edo and Hamada (2014). Morphological terminology of the adults generally follows Contreras-Ramos (1998), except wing venation and genital sclerites, which are based on Breitkreuz, Winterton, 300 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS and Engel (2017) and Liu et al. (2015a), respectively. The specimens reported in this work are housed in the following institutions:
ANDES-E Museo de Historia Natural, Universidad de los Andes, Bogot a, Colombia. CTNI Coleccion Taxonomica Nacional de Insectos ‘Luis Mar ıa Murillo’, Mosquera, Cundinamarca, Colombia. ICN Instituto Nacional de Ciencias Naturales, Universidad Nacional de Colombia, sede Bogot a, Colombia. MEFLG Museo Entomologico ‘Francisco Luis Gallego’, Universidad Nacional de Colombia, sede Medell ın, Colombia. MLS Museo de Historia Natural, Universidad de la Salle, Bogot a, Colombia. CAUD Museo de Historia Natural, Universidad Distrital Francisco Jos ede Caldas, Bogot a, Colombia. MUSENUV Museo Entomologico Universidad del Valle, Cali, Colombia. UDEA Museo Entomologico de la Universidad de Antioquia, Medell ın, Colombia. UNAB Museo Entomologico, Facultad de Agronom ıa, Universidad Nacional de Colombia, sede Bogot a, Colombia.
Additionally, the following museums are cited in the text:
MIZA Museo del Instituto de Zoolog ıa Agr ıcola ‘Francisco Fern andez Yepes’, Facultad de Agronom ıa, Universidad Central de Venezuela, Maracay, Venezuela. UMSP Department of Entomology, University of Minnesota, St. Paul, United States. ZMB Museum fur€ Naturkunde der Humboldt-Universit€at, Berlin, Germany.
Taxonomy Family Sialidae Sialidae currently comprises the Stenosialis lineage (Stenosialis þ Leptosialis) from Australia and South Africa, the Ilyobius lineage ((Ilyobius þ (Indosialis þ Haplosialis)) from Europe, Asia, India, Madagascar, and the Neotropics, and the Sialis lineage (Protosialis þ Sialis) of the Holarctic Region, Asia, and the Neotropics (Liu et al. 2015b). Adults are distinguished by the following characteristics: small size (forewing length 15 mm or less), ocelli are absent, prothorax wider than long and legs with the fourth tarsomere bilobed. Immatures are characterized by the presence of seven pairs of abdominal lateral filaments articulated, and the presence of a long, tapered, caudal filament on the last abdominal segment (New and Theischinger 1993; Liu et al. 2015b). The Neotropical fauna comprises two genera, Protosialis van der Weele, 1909 with a single species from Cuba, and Ilyobius with nine species distributed from AQUATIC INSECTS 301
Mexico to Argentina, of which two species are known from Colombia (Liu et al. 2015b, 2015c).
Genus Ilyobius Enderlein, 1910 Ilyobius Enderlein 1910: 381. Type species: Sialis flavicollis Enderlein, 1910: 380, by original designation.
The genus Ilyobius was erected by Enderlein (1910) from a single specimen from Tolima, Colombia. Nevertheless, Neotropical sialids were treated under the genus Protosialis for long time (Contreras-Ramos 2008; Liu et al. 2015b). Protosialis was proposed by van der Weele (1909) for three Neotropical species, (P. bifasciata (Hagen, 1861), P. chilensis (McLachlan, 1871), and P. mexicana (Banks, 1901)) plus a Nearctic species (Sialis americana (Rambur, 1842)), of which the latter was designated as the type species (Contreras-Ramos 2008; Liu et al. 2015b, 2015c). Later, Ilyobius was proposed as junior synonym of Protosialis by Penny (1981). Recently, in a com- prehensive phylogenetic analysis of the family, Liu et al. (2015b) recovered the genus as valid. These authors also assigned two additional species each to Protosialis (for a total of three) from North America and Cuba. Thus, Ilyobius comprises the vast majority of Neotropical sialids, with nine species distributed from Mexico south to Chile and Argentina (Liu et al. 2015b, 2015c; Archangelsky, Pessacq, and Berrondo 2017). Until now, Ilyobius flavicollis (Enderlein, 1910) and Ilyobius curvata Liu, Hayashi, and Yang, 2015 have been reported from Colombia.
Diagnosis of the adults Adults of this genus are medium sized insects (forewing length 7.0–17.0 mm in males, 9.0–19.5 mm in females), with head and pronotum generally orange to reddish brown, sometimes with dark spots; wings have RP with two branches. Male genitalia present endophallus internally with spinous and blackish setae, and gonocoxites þ gonapophy- ses 8 form a strongly sclerotized complex in the females (Liu et al. 2015b).
Biology The group is extremely rare in entomological collections, and its biology is virtually unknown. Contreras-Ramos (2008) hypothesizes that such rareness may be related to ecological factors such as low population densities and patchy distributions. Immature stages of I. flammata (Penny, 1981) and I. chilensis have been described (Azev^edo 2003, 2009; Archangelsky et al. 2017), but there is a formally published description only for the second. Species of this genus seem to be associated with small streams having low current velocity, soft and muddy substrates, with copious amounts of decomposing plant material, and they feed mainly on dipteran larvae and other small arthropods (Contreras-Ramos 2008; Azev^edo 2009; Clavier, Contreras- Ramos, and Guillemet 2009; Archangelsky et al. 2017).
Distribution Argentina, Brazil, Bolivia, Chile, Colombia, French Guiana, Mexico, Panama, and Venezuela (Liu et al. 2015b). 302 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 1. Habitus of Colombian species of Megaloptera: (a) Ilyobius curvata Liu, Hayashi, and Yang, 2015, holotype male (photo by Xingyue Liu); (b) Chloronia bogotana van der Weele, 1909; (c) Chloronia marthae Contreras-Ramos, 2002; (d) Corydalus affinis Burmeister, 1839; (e) Corydalus armatus Hagen, 1861; (f) Corydalus batesii McLachlan, 1868. Photos by Francisco Gamarra.
Ilyobius curvata Liu, Hayashi, and Yang, 2015 (Figure 1a) Ilyobius curvata Liu, Hayashi, and Yang 2015c: 56.
Type locality Colombia.
Diagnosis Forewing length, 7.0–8.6 mm in males, and 9.0–9.5 mm in females; head orange with dark brown spot around compound eye, medially bifurcate, and posteriorly trifurcate; Wings smoky brown, slightly darker on costal space, and proximal portion (Figure 1a). Gonocoxites 11 transversely arched proximad, gonostyli 11 very close to each other and strongly curved ventrad with unguiform tips (modified from Liu et al. 2015c).
Comments According to Liu et al. (2015b), this species is closely related to I. mexicana, based on colour pattern and genital morphology. Ilyobius curvata may be separated from I. AQUATIC INSECTS 303 mexicana by a strongly ventrally curved gonostyli 11 with pointed apex and a strongly narrowed posterior part of gonocoxites 8 of the female (Liu et al. 2015c). However, both species are similar to I. flammata from Brazil and Peru (Contreras- Ramos 2008), so further morphological and molecular work is necessary to delimit these species clearly. The distribution of I. curvata in Colombia is unknown, although the authors mapped the type locality towards the north, apparently in Antioquia, yet cited material does not specify a locality. Adults of I. curvata have been collected dur- ing February and May in Panama (Liu et al. 2015c). Larvae are unknown.
Distribution Colombia and Panama.
Ilyobius flavicollis (Enderlein, 1910) Sialis flavicollis Enderlein 1910: 380.
Type locality Colombia (Natagaima).
Diagnosis Forewing length 14.5 mm, hindwing length 13.0 mm. Head mainly blackish brown, dark orange in the medial region of vertex; antenna piceous; pronotum orange; wing membrane smoky brown, venation brown (modified from Liu et al. 2015c).
Comments The type specimen is in fragments and the abdomen is missing. The species was redescribed and illustrated by Liu et al. (2015c) from the fragments available, how- ever, fresh material needs to be collected, as the genital structures are unknown. The species is known solely from its type locality, the Natagaima Municipality, Tolima (Enderlein 1910). Immature stages are unknown.
Distribution Colombia.
Family Corydalidae Subfamily Corydalinae Adults of Corydalinae are recognized by a conspicuous postocular spine, male gono- stylus 9 well developed and clasper–shaped, and gonostyli 10 as small lobes. Larvae have ventral tracheal gills on the first eight abdominal segments and a pair of prolegs equipped with two claws on the last abdominal segment. This subfamily encompasses the Chloroniella lineage from South Africa (this genus with one species), the Protohermes lineage (Protohermes þ Neurhermes), the Neoneurmus lineage (Nevromus þ Neoneurmus), and the Acanthacorydalis lineage from Asia, as well as the Corydalus lineage (Chloronia þ Platyneuromus þ Corydalus) from the New World 304 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
(Contreras-Ramos 2011; Liu, Hayashi, Lavine, and Yang 2015d). The most recent phylogenetic hypothesis within the Corydalus lineage places Platyneuromus van der Weele, 1909 as sister group of Chloronia þ Corydalus. Meanwhile, the Asian genus Acanthacorydalis van der Weele, 1907 has been recently recovered as sister to the New World Corydalinae (Liu et al. 2015d). Corydalinae has the highest richness in the Neotropics, with 56 species grouped in three genera: Platyneuromus (3 spp.), endemic to Central America and Mexico; Corydalus (36 spp.), widely distributed but absent in the Caribbean Islands and Chile; and Chloronia (18 spp.), known from Mexico south to Brazil, Bolivia and Peru, including the Lesser Antilles (Contreras- Ramos 1998, 2005).
Genus Chloronia Banks, 1908 Chloronia Banks 1908: 30. Type species: Hermes corripiens Walker, 1860: 180, desig- nation by van der Weele (1909, p. 252).
The genus Chloronia comprises 18 species distributed from Mexico to Brazil, Bolivia, and Peru, including the Lesser Antilles (Dominica) (Contreras-Ramos 2005). The group was revised by Penny and Flint (1982), and subsequently several new species were described from Costa Rica (Flint 1992), Ecuador and Guatemala (Contreras- Ramos 1995), Brazil (Contreras-Ramos 2000), Venezuela (Contreras-Ramos 2002), and Bolivia and Peru (Contreras-Ramos 2006). Contreras-Ramos (2004a) also made a detailed redescription of C. bogotana van der Weele, 1909 because the identity of this species was repeatedly questioned due to the loss of the genitalia of the type specimen (Contreras-Ramos 2004a).
Diagnosis of the adults Adults of this genus are easily recognized by a body colour pattern of pale yellow with black spots; they are medium-sized insects (forewing length 24–50 mm), wings have the last branch of RP forked (Glorioso 1981; Contreras-Ramos 2002).
Diagnosis of the larvae Larvae of this genus are scarcely known. The larval head and pronotum are without muscle scars, and they often have two pairs of dark spots on the pronotum, which sometimes are barely noticeable. The latter character is not always diagnostic, as some South American species present the pronotum completely dark reddish brown, with two lateral darks stripes, or four short stripes instead dots. The larva of Chloronia hieroglyphica (Rambur, 1842) was described by Penny and Flint (1982) and Azev^edo (2003).
Biology Habitat and biology of Chloronia are little known. Larvae of C. hieroglyphica has been collected from small shaded streams with rocky or sandy substrates and an availability of plant material (Penny and Flint 1982; Geijskes 1984; Azev^edo 2009). Larvae of C. mexicana Stitz, 1914 have been found in submerged logs located in backwater zones AQUATIC INSECTS 305 of a river of seven meters wide and 2.5 m deep (Contreras-Ramos 1999b). An undetermined Mexican species of this genus has been found cohabiting with Corydalus larvae in a river 4 m deep (Contreras-Ramos 1999b).
Distribution From Mexico to Bolivia, Peru, and Brazil, including Caribbean Islands (Dominica).
Chloronia bogotana van der Weele, 1909 (Figure 1b) Chloronia bogotana van der Weele 1909: 253.
Type locality Colombia (Bogot a).
Material examined 1 $, Cundinamarca, San Francisco, San Miguel, 4 5803900N, 74 1703400W, 1570 m, 8.IV.2004, S. Florez, manual collection (UNAB).
Diagnosis Forewing length 41.1–47.2 mm; head with posterolateral broadened mark, anteriorly extended as a narrow stripe along suture limiting vertex. Forewing crossveins bor- dered with greyish spots; wing membrane with greyish spots on radial and medial cells, and posterior wing margin (Figure 1b). Fore tibia with a black spot at base. Gonostylus 9 incurved, fusiform with pointed apex; sternite 9 hexagonal, each lateral half triangular, posteromedial lobes not developed, lateral lobes well developed. Gonocoxites þ gonostyli 10 as a convex plate, with an anteromedial triangular projec- tion. Gonostyli 10 widely separated, papiliform, nearly two times as long as wide (Contreras-Ramos 2004a).
Comments This species is endemic to Colombia, and originally described from Bogot a. However, collecting data were probably erroneous, the species appears to be limited to eleva- tions below 2600 m a.s.l. The type specimen was probably from a locality close to Bogot a at a lower elevation and warmer climate. Until now, C. bogotana has been recorded from Cundinamarca (San Francisco Municipality), Cauca, and Meta (Villavicencio), and from altitudes of 2150 m a.s.l. or less (Penny and Flint 1982; Contreras-Ramos 2004a). Adults of this species have been captured in the months of March and August (Penny and Flint 1982; Contreras-Ramos 2004a). Immature stages remain unknown.
Distribution Colombia. 306 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Chloronia gloriosoi Penny and Flint, 1982 Chloronia gloriosoi Penny and Flint 1982: 11.
Type locality Panama (Chiriqu ı).
Material examined 1 #, Valle del Cauca, Bajo Anchicay a, 400 m, II.1985, leg. E. Burbano (MUSENUV).
Diagnosis Forewing length 39.0–45.0 mm in males, 42.0–44.0 mm in females; head generally with a long angular mark rarely divided into two spots, located on the posterior mar- gin of the occiput. Wings with basal and apical dark spots on the radial and medial cells; marginal cells with longitudinal medial dark stripes, especially on the anal and cubital areas. Fore tibia with a black spot basolaterally. Sternite 9 with anterolateral angles produced as trianguloid lobes, posterior margin having a pair of semi-triangu- lar lobes, submedially located. Gonostylus 9 incurved with a broadened basolateral angle, apical tooth displaced to the apical inner margin. Gonostyli 10 cylindrical, pap- illiform, nearly 2.5 times as long as wide, widely separated (Penny and Flint 1982).
Comments This species is known from montane rainy forests in Panama and Costa Rica; the only known altitudinal record is 1000 m a.s.l. (Flint 1992). Adults have been collected from April to mid-June (Penny and Flint 1982; Flint 1991). Herein, this species is recorded from Colombia for the first time, and from a single male specimen collected in lowland forests of the Anchicay a River basin (Valle del Cauca). The specimen was captured during February, at an elevation of 400 m a.s.l. Immature stages of this species are unknown.
Distribution Costa Rica, Colombia, and Panama.
Chloronia marthae Contreras-Ramos, 2002 (Figures 1c, 3a) Chloronia marthae Contreras-Ramos 2002: 59.
Type locality Venezuela (T achira).
Material examined 1 #, Caquet a, no further locality data (ANDES-E)
Diagnosis Forewing length 27.2–35.3 mm in males and 29.5–33.2 mm in females. Head with a pair of posterolateral ovoid blackish spots, anterior to occiput. Forewing with some AQUATIC INSECTS 307
Figure 2. Habitus of Colombian species of Megaloptera: (a) Corydalus clauseni Contreras-Ramos, 1998, holotype male (photo by Robin Thomson); (b) Corydalus colombianus Contreras-Ramos, 1998, holotype male (photo by Robin Thomson); (c) Corydalus flavicornis Stitz, 1914;(d)Corydalus muzoensis Ardila-Camacho, 2014, holotype male; (e) Corydalus nubilus Erichson, 1848; (f) Corydalus peruvianus Davis, 1903. Photos by Francisco Gamarra. greyish spots on membrane of radial and medial cells, with single or paired spots on cubital and anal spaces (Figure 1c); posterior margin at basal half of wing with numerous dark spots. Gonostylus 9 incurved, subclavate with apical point sclerotized, and a preapical tuft of long and thick setae. Sternite 9 with lateral and posteromedial lobes well developed. Gonocoxites 10 forming a convex plate, possessing strong anterolateral projections; gonostyli 10 sclerotized, triangular in shape, and strongly pointed (Contreras-Ramos 2002).
Comments Chloronia marthae is currently considered a possible junior synonym of the widely distributed Chloronia mirifica Nav as, 1925 (Contreras-Ramos 2005). These species were separated by the antennal length, the presence of a preapical tuft of setae on the gonostylus 9 and the pointed gonostyli 10 of C. marthae (Contreras-Ramos 2002). In subsequent publications, it was commented that there exist specimens from several locations of the Neotropical region identified as C. mirifica with an apparent variation regarding the presence of the tuft of setae of the gonostylus 9, and thereby, it is pos- sible that C. marthae may be part of a complex of geographical variation (Contreras- Ramos 2004b). In the present study, an individual from the Colombian Amazonia (Caquet a) was examined, and the genital structure matched the description of 308 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 3. Male genitalia of Colombian species of Megaloptera: (a) Chloronia marthae Contreras- Ramos, 1998;(b) Corydalus affinis Burmeister, 1839; (c) Corydalus armatus Hagen, 1861; (d) Corydalus batesii McLachlan, 1867; (e) Corydalus flavicornis Stitz, 1914; (f) Corydalus peruvianus Davis, 1903; (g) Corydalus nubilus Erichson, 1848; (h) Corydalus muzoensis Ardila-Camacho, 2014; (i) Corydalus liui sp. n. Scale bars: 1mm. C. marthae. Nonetheless, the number of flagellomeres (46), and the short antennal length is in accordance with C. mirifica. This specimen supports C. marthae as junior synonym of C. mirifica. A study of morphological and genetic variation of these spe- cies along their geographical range will be required to clarify the taxonomic status of C. marthae.
Distribution Colombia and Venezuela. AQUATIC INSECTS 309
Chloronia mirifica Navas, 1925 Chloronia mirifica Nav as 1925: 195. Chloronia hieratica Nav as 1928a: 11, 12; Penny and Flint, 1982: 10. Type locality Costa Rica (Cartago).
Material examined 1 $, Antioquia, Cocorn a, 1286 m, VIII.1956, F. Gallego, on shrubby vegetation, iden- tified by O. Flint (MEFLG).
Diagnosis Forewing length 29.0–40.0 mm in males and 25.0–42.0 mm in females; head with two brown spots on the posterior margin of occiput. Forewing with some radial cells, and all medial cells possessing dark spots basally and apically; posterior marginal cells with a single elongate dark spot. Gonostylus 9 incurved, apically inflated, with a strongly sclerotized apical point and a tuft on long setae. Sternite 9 with anterolateral angles produced, and with an elongate lobe in the posterior margin. Gonostylus 10 elongate, triangular, as long or longer than wide, ending in an apical point (Penny and Flint 1982).
Comments This Chloronia species is probably the most common and widely distributed in the Neotropics. Collecting records have been reported in the warmer months of the year from May and June and from August to October in Mexico and Guatemala. In Costa Rica and Panama, there are records from January, and from March to November; in Colombia and Ecuador, there are records from December to March and May (Penny and Flint 1982; Flint 1992; Contreras-Ramos 1995; 2004b). In Colombia, this species has been reported solely from Meta department (Villavicencio), at about 450 m a.s.l. (Penny and Flint 1982). Immatures of C. mirifica are unknown.
Distribution Colombia, Costa Rica, Ecuador, Guatemala, M exico, Panama, and Peru.
Genus Corydalus Latreille, 1802 Corydalus Latreille 1802: 290. Type species: Hemerobius cornutus Linnaeus, 1758: 551, by monotypy. Corydalis Latreille 1804: 44; ‘Lapsus calami’, Glorioso 1981: 284. Corydalium Oken 1815: 856; Oswald and Penny 1991: 61. Corydalia Billberg 1820: 95; Oswald and Penny 1991: 61. The genus Corydalus is the most diverse, widely distributed, and best known of the New World megalopterans. Representatives in this group are known from southern Canada to Argentina, with the noteworthy exception of Chile and the Caribbean Islands (Contreras-Ramos 1999a, 2005). Contreras-Ramos (1998) carried out a 310 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS taxonomic revision and phylogenetic analysis based on morphological characters. Subsequently, new species from Venezuela and Colombia were described (Contreras- Ramos 2002; Contreras-Ramos and von der Dunk 2010; Ardila-Camacho 2014). The phylogeny of the genus was recently updated to include the Venezuelan species described up through 2002 (Contreras-Ramos 2011). Until now, the genus includes 36 species, of which 31 are Neotropical in distribution.
Diagnosis of adults Adults of Corydalus are recognized from other Neotropical dobsonflies by their max- illary and labial palpi with four and three palpomeres, respectively; although there are two South American species with a partial subdivision in the last palpomere of both (Contreras-Ramos 1998). The genus is highly heterogeneous in size, with a forewing length varying from 33.5 to 85.3 mm in males, and such variation often plays a part in intraspecific phenotypic plasticity as well. Body colour pattern is generally of pale brown to greyish brown and dark reddish brown; wings are semi-translucent, with a colour pattern of a pale brown, greyish brown or reddish brown background, often with fenestrate pale areas, dark suffusions or spots, frequently small semicircular fen- estrate white spots, and sometimes with pale pterostigma-like areas. Several species have a marked sexual dimorphism, in which the male mandibles are hypertrophied with partly or completely reduced dentition. The size of the mandibles has a positive allometric pattern in function of the body size; this trait is a derived character within the group, as in the most basal species both sexes have monomorphic mandibles. It is known that this trait is involved in sexual selection, and independently evolved in Acanthacorydalis and Corydalus. Also in Corydalinae, there exists a type of ornamen- tation in the genus Platyneuromus that consists in an exaggerated development of the male postocular flanges (Liu et al. 2015d; Ram ırez-Ponce, Garfias-Lozano, and Contreras-Ramos 2017).
Diagnosis of the larvae Larval stages are similar to those of Platyneuromus, in that both have the head and thoracic nota with muscle attachment scars. According to Contreras-Ramos and Harris (1998), larvae from both genera may be separated by abdominal setal charac- ters. For example, in Corydalus there are erect dark macrosetae, which are striated, inflated, and apparently closed at the apex, or elongated, striated or completely smooth. In contrast, Platyneuromus has clear setae, which are either tubular and wid- ened with an open apical portion, or thin, smooth and with an inconspicuous apical hole. Furthermore, Platyneuromus larvae present long and pointed submental projec- tions that surpass the anterior margin of the mental plates. Many Corydalus species have short and blunt submental projections, but some South American species have these projections similar to those of Platyneuromus (Contreras-Ramos and Harris 1998; Azev^edo and Hamada 2007, 2007). Hitherto, detailed descriptions of the last larval instar (Azev^edo and Hamada 2006, 2007) have been made for three Neotropical species of Corydalus. Other larval descriptions are available in Azev^edo (2003, 2009) and a key to species and genera for Brazilian species was recently published by Azev^edo and Hamada (2014). AQUATIC INSECTS 311
Biology Immatures of Corydalus are voracious predators feeding on a wide variety of aquatic invertebrates, or even small aquatic vertebrates, but they are also known to be scav- engers and cannibalistic (Contreras-Ramos 1998). Larvae are found in lotic ecosys- tems, including small streams and creeks and large rivers, generally with cold and well oxygenated waters (Geijskes 1984). Many species are generalists regarding niche preference, and may be collected from beneath rocks, logs or decomposing sub- merged plant material and woody debris. Sometimes several species have been found that appear to be cohabiting in a same habitat (Ardila-Camacho, personal observa- tion). Larval stages encompass the majority of the life cycle; they undergo around 11 molts before leaving the water to pupate. Adults have a life-span of around one to two weeks depending on the sex, and are attracted to light (Contreras-Ramos 1999a; Azev^edo and Hamada 2007).
Distribution From South Canada to Argentina, except Caribbean Islands and Chile.
Corydalus affinis Burmeister, 1839 (Figures 1d, 3b) Corydalus affinis Burmeister 1839: 951.
Type locality South America, Brazil(?).
Material examined Caldas. 1 $, La Dorada, 25.IV.1992, leg. M. Garc ıa (UNAB).
Cundinamarca. 1 $, Anolaima, 14.IV.1992, leg. G. Gomez (UNAB).
Meta. 1 #, Macarena, casco urbano, 3 380N, 73 540W, 443 m, 28.VIII.2016, leg. G. Jimenez (UNAB).
Tolima. 1 $, Espinal, 6.IV, leg. R. Espinel (UNAB). 1 #, Mariquita, casco urbano, 5 1204900N, 75 104500W, 465 m, 16.V.2011, leg. J. Sanchez, manual collection (UNAB). 1 $, Saldana,~ 14.IV.1993, leg. J. Barriga (UNAB). 1 #, Armero, III.1964, leg. Mar ın (CTNI). 1 #, Honda, 9.IX.1964, leg. J. Delgado (CTNI). Valle del Cauca: 2 $, Palmira, V.1943, leg. B. Lozada (CTNI); 1 $, Palmira, 6.VI.1961, leg. L. Agudelo (CTNI).
Diagnosis Forewing length 32.0–58.0 mm in males, 38.6–61.6 mm in females. Head flattened with subtle postocular dark stripe; male mandibles generally elongate with reduced dentition; filiform antennae, slender, very dark brown to black. Forewing pale brown with RA space, proximal cells of RP and medial cells with a series of pale areas 312 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS surrounded by brown suffusions (Figure 1d). Male genitalia with ectoproct elongate, flattened at apical 1=4, densely covered by fine setae; gonostylus 9 similar to ectoproct (Figure 3b); sternite 9 subquadrate; gonostyli 10 subquadrate, nearly two times as long as wide, and moderately convergent (Contreras-Ramos 1998).
Comments This species is commonly found in the lowlands of the Amazon basin, and the foot- hills of the mountainous regions of Colombia and Ecuador. Altitudinal distribution encompasses from 54 to 600 m a.s.l. In Colombia, it is known from Antioquia, Boyac a, Cundinamarca, and Tolima in the Andean region; from Choco in the Pacific region, and from Arauca, Casanare, and Meta in the Orinoqu ıa (Contreras-Ramos 1998; Ardila-Camacho 2014). The last larval instar was described by Azev^edo (2009).
Distribution Argentina, Bolivia, Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, and Venezuela.
Corydalus armatus Hagen, 1861 (Figures 1e, 3c, 4, 5) Corydalis armata Hagen 1861: 321. Corydalus quadrispinosus Stitz 1914: 197; Contreras-Ramos 1998: 48.
Type locality Venezuela (no further locality data).
Material examined, adults Antioquia. 1 #, Versalles, Fca. Los Naranjos, 5 5506000N, 75 340000W, 1800 m, 12.IV.2012, leg. G. Villamizar, entomological net (UNAB). 1 #, Medell ın, 14.VII.2008, El Poblado, leg. A. Gulfo, manual collection (UDEA). 1 #, same data but cerro El Picacho, VI.1984, leg. L. Osorio, manual collection (UDEA). 1 #, same data but Barrio Manrrique, 7.VII.2001, leg. N. Herrera, manual collection (UDEA). 1 #, Itagui,€ Barrio Pedregal, 26.XI.2005, leg. E. Escobar, manual collection (UDEA). 1 #,El Penol,~ IX.1987, leg. A. Arizmendi, manual collection in flight (UDEA). 1 #, Yarumal, finca La Mar ıa, 26.II.2012, no collector, manual collection (UDEA-63268). 1 #, Argelia, 5 4403300N, 75 804700W, 1750 m, leg. J. C. Castrillon, manual collection in for- est (UDEA-63035). 1 #, Maceo, corregimiento Puerto Nus, vereda San Pedro, finca San Pedro, 6 270N, 74 470W, 890–1100 m, 17–19.II.2012, no collector, manual collec- tion (UDEA-62660).
Boyaca. 2 #, Sutatenza, 20.III.1973, leg. Hno. Alfonzo Juan (MLS-006, 007).
Caldas. 1 #, Norcasia, Moscovita, El Encanto, 74 5203300N, 5 3402000W, 600 m, leg. D. Quintana, entomological net (UNAB). 1 #,P acara, 17.IV.2000, no collector (UNAB). AQUATIC INSECTS 313
Figure 4. Last instar larva of Corydalus armatus Hagen, 1861: (a) habitus, dorsal; (b) habitus, ventral (photos by Francisco Gamarra); (c) head, dorsal; (d) head, ventral; (e) thorax, dorsal; (f) forefemur, dorsal; (g) mandible; (h) spiracle of abdominal segment 1; (i) sternellum, lateral; (j) last two abdom- inal segments, dorsal; (k) macro- and microsetae of abdominal tergite 5; (l) V-shaped pattern of abdominal sternite 5.
1 #, Alto de San Miguel, vereda La Clara, 6 202700N, 75 3808100W, 1900 m, 10.X.2005, no collector, manual collection near stream margin (UDEA).
Cauca. 1 #, Popay an, Urbanizacion La Virginia, 2 270N, 76 370W, 1760 m, 4.VIII.2011, leg. C. Mera, manual collection (UNAB). 314 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 5. Pupa of Corydalus armatus Hagen, 1861: (a) exuvia of an emerged adult from a pupal chamber observed below of a large stone close to a water stream in Chipat a, Santander; (b) pupa in the pupal chamber observed behind the glass of a beaker in the laboratory, see the prepupa exuvia near the abdomen; (c, d), male genital primordia: (c) dorsal; (d) ventral. Scale bars: 1 mm.
Cundinamarca. 1 #, La Mesa, Vda. Calucata, Fca. Frutales, 4 380N, 74 280W, 1200 m, 4.XII.2012, leg. D, Tovar, manual collection (UNAB). 1 #, Anapoima, Vda. Las Mercedes, 680 m, 29.III.2010, leg C. Rodriguez, entomological net (UNAB). 1 #, Pacho, V.1999, leg. A. Vergara (UNAB). 1 #, La Calera, Vda. Quisquisa, 25.VIII.1998, leg. L. Avila, F. Ardila (UNAB). 1 #, San Francisco, 4 5803900N, 74 1703400W, 1570 m, 10.IV.2004, leg. C. Calderon, entomological net (UNAB). 1 #, La Mesa, 5.IV.1994, leg. H. Rodr ıguez (UNAB). 1 #, Tabio, 2.XII.1995, no collector (UNAB). 1 #, Quipile, La Sierra, 7.I.1996, leg. G. Parra (UNAB). 1 #, Gachet a, casco urbano, 4 490100N, 73 3705900W, 1780 m, 18.X.2011, leg. F. Torres, manual collection (UNAB). 1 #, Sasaima, Santa Ana, 31.I.2008, leg. V. Bernal, K. Turriago (UNAB). 1 #,El Colegio, 31.V.1996, leg. J. Ardila (UNAB). 1 #, San Antonio del Tequendama, Vda. San Jos e, 4 420700N, 74 1303100W, 1463 m, 18.IV.2015, leg. A. Garc ıa, entomological net (UNAB). 1 #, Pacho, 5 80N, 74 90W, 2136 m, 28.II.2015, leg. L. Zamora, manual collection (UNAB). 1 $, Sasaima, no collector (MLS-15). 1 #, Chocont a, 9.XI.1980, leg. L. Cort es (MLS-10).
Huila. 1 #, Palestina, Vda. Para ıso, Fca. El Vergel, 1 440N, 76 90W, 2.IV.2014, leg. L. Pardo, manual collection (UNAB). AQUATIC INSECTS 315
Meta. 1 #, Villavicencio, 4 90N, 73 380W, 467 m, 11.VII.2013, leg. G. Beltr an, manual collection on Psidium guajava (UNAB). 1 #, Villavicencio, 4 80N, 73 370W, 467 m, 8.X.2013, leg. J. Jim enez, entomological net (UNAB). 1 #, Villavicencio, Vda. Bueno Aires, Fca. La Esmeralda, 4 70N, 73 390W, 558 m, 17.XI.2014, leg. S. Rodr ıguez, man- ual collection (UNAB). 1 #, Granada, casco urbano, 3 3204000N, 73 4202300W, 332 m, 9.IV.2012, leg. Y. Mar ın, manual collection (UNAB). 1 #, Puerto Lopez, Llano Grande, 4 50N, 72 570W, 178 m, 17.V.2005, leg. E. Romero, entomological net (UNAB).
Narino.~ 1 #, Ricaurte, Balneario el Para ıso, 1 540N, 78 40W, 1810 m, 15.III.2012, leg. H. Benavides, entomological net (UNAB). 1 #, Linares, Vda. El Motilon, 1 230N, 77 320W, 2109 m, leg. J. Paredes, manual collection (UNAB).
Risaralda. 1 #, Santa Rosa de Cabal, Fca. El Ensueno,~ 4 410N, 75 370W, 7.V.2013, leg. A. Guataqu ı, manual collection (UNAB). 1 #, Santa Rosa de Cabal, 4 520N, 75 370W, 1715 m, 6.IV.2012, leg. M. Ram ırez, manual collection (UNAB).
Santander. 1 #, Parque Nacional Chicamocha, 6 450N, 73 20W, 1200 m, 15.XI.2013, leg. A. Guataqu ı, manual collection (UNAB). 1 #, Barbosa, 15.V.1969, no collector (UNAB). 1 #, La Belleza, Vda. Zeilon, fca. Los Arrayanes, 5 510N, 73 580W, 2050 m, 15.X.2012, leg. S. Torres, entomological net (UNAB). 1 $, Chipat a, Quebrada Roperana, 30.VIII.2015 as mature larva, pupa: 4–16.XI.2015, leg. A. Ardila (CAUD).
Tolima. 1 #, Purificacion, 5.XI.1995, leg. D. Mora (UNAB). 1 $, Melgar, 28.XI.1976, Hno. Niceforo (MLS-22).
Valle del Cauca. 1 #, Tulu a, 1.X.2003, 800 m, leg. O. Combita, manual collec- tion (UNAB).
Material examined, larvae 16 larvae (L), Santander, Chipat a, Roperana stream, 6 0304500N, 73 3802400W, 1700 m, 30.VIII.2015, leg. A. Ardila, (CAUD). 20 L, same data but 28.VI.2015, leg. A. Ardila (CAUD). 20 L, same data but 28.XII.2014, leg. A. Ardila (CAUD). 10 L, same data but 25.IX.2015, leg. A. Ardila (CAUD). 10 L, same data but 18.XII.2015, leg. A. Ardila (CAUD).
Material examined, pupa 1 pupa, then $,4–16.XI.2015, reared from a mature larva, Chipat a, Quebrada Roperana, 30.VIII.2015, leg. A. Ardila (CAUD).
Diagnosis, adult Forewing length 44.7–82.15 mm in males, 51.0–79.0 mm in females. Male mandibles elongate with a positive allometric pattern as a function of body size, with reduced dentition. Male antennae thickened, slightly subserrate, pale to dark brown, including scape and pedicel. Forewing nearly uniformly pale brown, with small, semicircular, 316 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS fenestrate white spots (Figure 1e). Male genitalia with tergite 9 subquadrate; ectoproct elongate, tubular with an apical incurvation; gonostylus 9 elongate, subclavate; fused gonocoxites 10 forming a convex plate, with long anterolateral projections; gonostyli 10 papilliform, usually as long as wide, with less than half length surpassing the pos- terior margin of the gonocoxites 10, slightly divergent (Figure 3c); conspicuous and well developed pregenital sacs (Contreras-Ramos 1998). This is a widely distributed montane species found throughout Andean mountain ranges. Corydalus armatus is morphologically very similar to C. peruvianus and C. clauseni, although the three species can be easily separated by the colour pattern of the antennae and wings, and the morphology of the male genital structures. Conversely, the females are quite similar, and do not have a clear morphological delimitation. Even the genital structures are virtually indistinguishable, so generally valid records of C. armatus are from male specimens.
Diagnosis, larva This species has a variable body colour pattern, in which the head presents a wide range of brown tones. Head of specimens range from uniformly very pale brown to a very dark brown patterned with a conspicuous pale brown area between frontal scler- ite and stemmata. The shape of the head is also variable; some specimens have a robust subquadrate head, while other have a relatively narrow head. The mandible has the basal tooth bluntly rounded and short, not fused to the third preapical tooth, and closer to the apex of the mandible than the base; it projects at an angle of 45 to the longitudinal axis of the mandible. The third preapical tooth is broad, short and rounded. Submental projections are short, blunt and not surpassing anterior margin of mental plates. The sternellum is short, bluntly rounded and covered with short and thick setae. Abdominal terga have very dark brown decumbent, star-shaped microsetae, and subglobose and striated dark macrosetae.
Description of last instar larva The following larval description is based mainly on mature larvae (> 70 mm by body length), however 55 larvae from different sizes (22.50–82.60 mm by body length) were also examined (Figure 4).
Head. Labrum subtrapezoidal, with anterior margin rounded and some amber setae, anteriorly very pale brown, medially with a transverse brown stripe, lateral margins dark brown, remainder of surface tan. Clypeus subrectangular, with medial region tan or pale brown, laterally brown, posterior margin tan. Mandible dark reddish brown, with five teeth, the apical and first preapical teeth long and pointed; third preapical tooth short, broad, and rounded, fourth one short and reduced; basal tooth situated at an apical 1/3 of mandible length, not fused to the fourth preapical tooth and form- ing an angle of 45 to the longitudinal mandible axis (Figure 4g). Gena dark brown on the margins, tan at center; genal carina dark brown. Frontal sclerite dark brown at anterior margin, laterally with two elongate pale brown spots, central region punctate, dark brown, laterally with a long seta, posterior region with pale brown scars. Antenna with five antennomeres, the first one dark brown, remainder antennomeres AQUATIC INSECTS 317 amber. Region between stemmata and frontal sclerite tan to pale brown, bordered with dark brown (Figure 4c). Frontal and coronal sutures bordered with brown. Ocular region with six stemmata arranged as a pentagon, with a single central stemma; adjacent surface to corneal lenses dark brown, with some long setae. Postocular region punctate, dark brown; adjacent region to postocular carina mottled, with a mixture of yellow and brown, entire surface with brown setae; postocular car- ina dark brown with a row of short, thick, pale brown setae. Region between postocu- lar carina and coronal suture punctate, dark brown, with muscle scars on the central part of head (Figure 4c). Occipital suture and carina dark brown on anterior margin, tan on the posterior margin. Occipital suture very dark brown; occiput predominantly very dark brown, laterally with a diffuse yellow spot, medially with a broadened yel- low spot. Cardo predominantly brown, posterolaterally pale brown, inner anterior corner with a tuft of setae; stipes ventrally mainly amber, except apex tan; entire sur- face densely covered by short and thick pale brown setae, inner margin with a row of long bristles; dorsal surface dark amber at margins, apex tan, remaining surface pale amber, glabrous, smooth and shiny; maxillary palpi with five palpomeres and pale amber in colour; galea pale amber. Labium with ligula tan, densely covered by short and fine setae; prementum tan or pale brown, dark brown on margins; mental plates with a mottled pigmentation pattern, consisting in a mixture of yellow and brown; submental projections short, blunt, not surpassing anterior margin of mental plates, with brown margins (Figure 4d); gula mottled, with a mixture of yellow and pale- to dark brown, and a pair of ellipsoid scars at the center, laterally with two pairs of long setae.
Thorax. Pronotum subquadrate, nearly as long as wide, with pale brown muscle scars in a punctate dark brown background (Figure 4e), margins dark brown, entire surface with scattered macrosetae as follows: long and tapering, short and thickened, and subglobose, striated, and dark, mainly at margins. Prosternum predominantly yellow, laterally with dark brown suffusions and a pair of long and narrow pale brown stripes, entire surface with scattered spinous setae; sternellum yellow, slightly pro- duced, bluntly rounded, in ventrolateral view, covered with short and thick setae (Figure 4i). Pleural membrane yellow, with numerous, subglobose, black macrosetae. Meso- and metanotum similar in shape and size, subrectangular, predominantly very dark brown, with pale yellow scars (Figure 4e), entire surface with at least four types of macrosetae: long, smooth, tapering, and brown; black, long and subclavate, mainly in the central region; black, subglobose, and striated; and black, tubular, and short. Meso- and metasternum and pleural regions yellow, with abundant, decumbent, star- shaped, dark microsetae, and short, subglobose, black macrosetae.
Legs. Coxae basally brown, pale amber at apex, entire surface covered by short spin- ous setae dark brown to black, on mid- and hind leg with some, subglobose, striated, black macrosetae. Trochanter yellow, with numerous, short, thick, spinous, pale amber setae. Femora mainly pale brown, dorsal surface with longitudinal narrow brown stripes and a row of hair-like setae, reminder surface covered by short spinous setae (Figure 4f), and long and tapering amber setae, mainly on ventral surface. Tibiae, dorsally infuscate with short spinous setae, ventral and lateral surfaces pale 318 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS brown with thick long tapering setae. Tarsi brown, ventrally with two rows of short setae, and numerous long, thin setae at apex; tarsal claws amber.
Abdomen. Terga yellow or tan, covered with star-shaped and widened microsetae, and subglobose and striated macrosetae, apparently closed at apex (Figure 4k). Spiracles reniform, dark brown (Figure 4h); a row of hair–like setae between the spir- acle and the lateral filament of each abdominal segment present. Lateral filaments tapering, thin (Figures 4a, b), dorsally with a thick row of hair-like setae, remainder of surface covered by subglobose black macrosetae, and striated, broadened and spin- ous microsetae, ventrally only microsetae present. Sternites densely covered by broad- ened, star–shaped microsetae on central region, and tubular, striated and apically widened, plus decumbent striated microsetae on the lateral parts. Sternites 2–8 lat- erally on each side with a pair of long tapering amber setae; sternites 2–6 with a glab- rous V-shaped area (Figure 4l). Sternites 9 and 10 sclerotized, with dense brush of brown setae. Cerci and prolegs densely covered by striated, decumbent and black, spinous and clear microsetae, and long and tapering, and subglobose striated black macrosetae (Figure 4j); claws amber.
Description of pupa Body length, 59.50 mm (n ¼ 1); head subquadrate, postocular flange and spine devel- oped, antenna short and coiled, pale brown; mandibles developed, slightly shorter than head, mainly brown, apically dark brown; compound eyes greyish brown, poorly developed. Prothorax subcylindrical, elongate, pale brown, but darkening with age. Pterothorax tan to pale brown; wing pads pale- to dark brown. Legs pale- to dark brown, base of tibiae slightly darker. Abdomen, soft, coiled, tan to pale brown, ter- gites with brown suffusions, entire surface covered with fine, pale brown setae, spi- racles brown. Male genital primordia with tergite 9 subquadrate in dorsal view, ectoproct short, lobe–shaped, bluntly rounded, densely setose (Figure 5c); gonostylus 9 shorter than ectoproct, lobe shaped. Fused gonocoxites þ gonostyli 10 plate shaped, ovoid, medially sub-divided, gonostyli 10 short lobe–shaped (Figure 5d).
Bionomics The specimens studied were collected in a small stream located in mountainous areas (over 1700 m a.s.l.) of Chipat a Municipality, Santander Department. The lar- vae were found beneath stones and logs, or among decomposing leaves, and sub- merged roots in riffles or pools located in shaded areas with riparian arboreal vegetation as well as in open areas with full sun. In the same stream, larvae of Corydalus liui were collected, but generally in different microhabitats, although sometimes both species were found in the same places. Furthermore, in the same region, but at a higher elevation (over 2012 m a.s.l.), adults of C. peruvianus Davis, 1903 were frequently collected. Interestingly, at that elevation no adults or larvae of C. armatus were found. Late instar larvae and prepupae were frequently collected during the dry season (December–February). Conversely, these stages were rarely collected in rainy seasons. However, according to Contreras-Ramos (1998), adults of this species have been AQUATIC INSECTS 319 reported throughout the year. During the dry season, numerous larvae were seen dur- ing early hours in the morning in shallow waters exposed to sunlight. On the other hand, they are very active during the night, and have been observed crawling in the bottom of the stream, when their stemmata reflect projected artificial light. During daytime, several larvae of different stages were observed, often very close one to the other and beneath the same stones. Pupae and exuviae were found in moist soil below large stones close to the stream (Figure 5a); sometimes prepupae and pupae of C. liui, were found in neighbouring places, although they were observed in very dry soil. In the laboratory, C. armatus larvae were fed every three days, and different behav- iours against different types of prey were observed. When earthworms (Eisenia fetida (Savigny, 1826)) were offered as food, the larvae struck the worm using their mandi- bles, and manipulated the prey using their prolegs; if the worm made violent move- ments to escape, the larva gave strong bites, and manipulated the prey searching one of its extremes with the aid of the mandibles and prolegs. Subsequently, the larva swallowed the worm with the aid of its maxillae and movements of the head, con- suming the prey without biting. Sometimes, the worm was larger than the larva, and even so it was completely devoured. When an insect, such as a cricket (Acheta domes- ticus (Linnaeus, 1758)), was offered, the larva caught the prey with a strong and rapid bite. Then, the insect was firmly grasped by the mandibles, and if the prey made vio- lent movements, the larva gave strong bites, until the insect was immobilized. Later, the prey was manipulated with the mandibles and prolegs until the head was found, which the larva cut and swallowed immediately after manipulating it using their man- dibles and maxillae while the prey body was held with the prolegs. It was observed that a late instar larva consumed an adult female of Acheta domesticus completely within 20 minutes. The feeding behaviour shown for swimming prey such as small fishes (Poecilia sp.) was not always successful. It was observed that the larva perceived a slight water disturbance when a fish swam nearby. When that happened, the larva made a rapid attack, even following the escape directions of the prey and sometimes partially leaving its burrow. Once the fish was caught, the larva squeezed it with its mandibles until the prey was disabled or killed. Subsequently, the larva manipulated the fish with its prolegs and mandibles while searching for an end. If the prey was large, the larva cut small pieces that swallowed with the aid of its maxillae. Conversely, if the fish was small the larva swallowed it completely without biting. After the prey was consumed, the larva proceeded to clean its head and mouth- parts using the prolegs. Sometimes, the struggle with the prey caused the burrow of the larva to become disturbed, and it was observed that the larva would move small stones and sand using its mandibles in order to reconstruct its burrow. When late instar larvae were induced to pupate in a container with a slightly mois- tened substrate, we observed the time used to construct the pupal chamber ranged from 20 days to two months. The larva constructed a semicircular or ellipsoidal chamber beneath a stone using its head and abdomen. Later, the larva adopted a ven- trally recurved position, and after a few days the pupa emerged (Figure 5b). It was observed that the emergence time of the adults varies with the sex, as female speci- mens took 12–14 days, whereas males took 17–19 days. 320 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Distribution Argentina, Bolivia, Colombia, Ecuador, Peru, and Venezuela. This species previously was known to occur in the Colombian departments of Boyac a, Cundinamarca, and Valle del Cauca (Contreras-Ramos 1998). Ardila-Camacho (2014) provided records from Atl antico, Antioquia, Caldas, Casanare, Guajira, Meta, Narino,~ Quind ıo, Santander, and Tolima. However, the record from Atl antico is here considered erro- neous. Label data from the specimen indicated it was collected from the ‘Sabanalarga’ Municipality, but the department was not mentioned. In Colombia there are three departments with a locality known with that name, so the specific provenance of the specimen is uncertain. Furthermore, Contreras-Ramos (1998) mentioned that this species is probably absent in northern Colombia. Altitudinal collecting records of C. armatus are comprised between 500 m and 2600 m a.s.l. (Contreras-Ramos 1998, 2004b).
Corydalus batesii MacLachlan, 1868 (Figures 1f, 3d) Corydalus batesii MacLachlan 1868: 232, 233.
Type locality Brazil (Amazonas).
Material examined Amazonas. 1 #, Finca El Rebano,~ 21.V.2014, leg. J. Melgarejo (ANDES-E).
Cundinamarca. 1 #, La Mesa, 30.I.1993, leg. A. Bonilla (UNAB).
Meta. 1 #, San Mart ın, Reserva Las Unamas, 3 540N, 37 110W, 240 m, 11.IV.2011, leg. L. Ojeda, light trap (UNAB). 1 $, Puerto Gait an, 5.IV.1994, leg. H. Roa (UNAB). 1 #, Villavicencio, 4.XI.2001, 700 m, leg. M. Gallen (UNAB).
Diagnosis Forewing length 45.85–64.0 mm in males and 52.0–67.6 mm in females. Head and pronotum dark reddish brown. Male mandibles elongate with reduced dentition, with a positive allometric pattern as a function of the body size. Filiform antennae, yellow with the apical 1/4 infuscate. Forewing with a contrasting colour pattern, costal cross- veins blackish; costal, subcostal, and anterior radial spaces infuscated; ram1, mcu1, and area surrounding M fork dark brown (Figure 1f). Male genitalia with tubular ectoproct, slightly narrower in the preapical region, apex dorsoventrally broadened, but without an incurvation (Figure 3d); sternite 9 subquadrate with posterolateral lobes slightly developed; gonostylus 9 elongate, subclavate; fused gonocoxites 10 form- ing a convex plate with a wide and short anteromedian projection; gonostyli 10 papil- liform, 1.5–2 times as long as wide, moderately mesally directed to nearly straight (Contreras-Ramos 1998). AQUATIC INSECTS 321
The last larval instar is approximately 7.30 cm in length, and it is distinguished by the region between stemmata, genal carina and medial region of frontal suture having a wide, pale yellow stripe. The mandible has the basal most tooth widened, and par- tially fused to the third preapical tooth. Abdominal terga with subtriangular black microsetae, and claviform, tubular or star-shaped macrosetae, pale- to dark brown (Azev^edo and Hamada 2007).
Comments This species has been reported to occur in the central and oriental cordilleras of Colombia, specifically in the departments of Antioquia, and Cundinamarca, respect- ively. It is a common species in the Orinoco and Amazon basins of Colombia as well, with records in the departments of Meta and Amazonas, respectively (Contreras- Ramos 1998; Ardila-Camacho 2014). The last larval instar and aspects of its biology were described in detail by Azev^edo and Hamada (2007). In addition, information about the ecology of the larvae is available in Azev^edo (2009).
Distribution This species is widely distributed in the Amazon basin and adjacent regions, includ- ing records from Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, and Venezuela (Contreras-Ramos 1998).
Corydalus clauseni Contreras-Ramos, 1998 (Figure 2a) Corydalus clauseni Contreras-Ramos 1998: 75.
Type locality Ecuador (Canar).~
Material examined Quind ıo. 1 $, Montenegro, 4 560N, 75 750W, 1620 m, 24.II.2012, leg. C. Pinilla, ento- mological net (UNAB).
Diagnosis, adult Forewing length 49.85–67.4 mm in males and 43.0–59.45 mm in females. Head pale- to dark brown, with a subtle postocular dark stripe. Male mandible elongated, with reduced dentition, with a positive allometric pattern in function of the body size. Antennae black in both sexes, sinuous and shiny in males. Forewing generally dark, golden brown, semitranslucent, subtly maculated with small, semicircular, fenestrate, white spots (Figure 2a). Male genitalia with elongated and tubular ectoproct, with a well-developed apical incurvation; gonostylus 9 elongated, subglobose; fused gonocox- ites 10 forming a convex plate; gonostyli 10 short, papilliform; adjacent area to gono- coxites 10 with very long setae; pregenital sacks well developed (Contreras- Ramos 1998). 322 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Comments Previously reported from Colombia in the lowland rainy forests of the Anchicay a River Basin and Buenaventura (Valle del Cauca) and Caldas departments (Contreras- Ramos 1998; Ardila-Camacho 2014). Altitudinal records are between 305 to 1460 m a.s.l., with the majority of them below 800 mm (Contreras-Ramos 1998). Adults have been collected in December, January, March, May, August, and October. Immature stages are unknown.
Distribution Costa Rica, Colombia, and Ecuador.
Corydalus colombianus Contreras-Ramos, 1998 (Figure 2b) Corydalus colombianus Contreras-Ramos 1998: 79.
Type locality Colombia (Valle del Cauca).
Diagnosis Forewing length 46.5–47.9 m in males, and 48.6–54.0 mm in females. Head dark red- dish brown, with slight muscle scars. Male mandible elongated with discrete dentition and a developed median protrusion, located at inner margin. Female mandible black with a slight basal protrusion. Filiform antennae, pale reddish brown with an apical 1/4 or 1/5 infuscate. Forewing of both sexes dark golden brown, strongly maculate with small to moderately large irregular pale areas surrounded by a dark brown suffu- sion (Figure 2b). Male genitalia with elongate and narrow ectoproct, with a narrow and digitiform apex; gonostylus 9 elongate, narrow, subcylindrical, slightly longer than ectoproct, equipped with a small protuberance at base and an apical small claw; fused gonocoxites 10 forming a convex plate, mesally narrowed, and with anterolat- eral projection poorly developed and pointed; gonostyli 10 widened, subquadrate, widely separated, divergent (Contreras-Ramos 1998).
Comments Corydalus colombianus is an uncommon species, and is known solely from the low- land rainy forests of the Anchicay a and Calima basins in the Valle del Cauca Department (Contreras-Ramos 1998; Ardila-Camacho 2014). The altitudinal distribu- tion range of this species is between 300 and 500 m a.s.l. Adults have been collected in the months of February and March, and July and August (Contreras-Ramos 1998). The larvae of this species are unknown.
Distribution Hitherto only known from and apparently endemic to Colombia. AQUATIC INSECTS 323
Corydalus flavicornis Stitz, 1914 (Figures 2c, 3e) Corydalus flavicornis Stitz 1914: 195. Corydalus nevermanni Nav as 1934: 507, 508; Contreras-Ramos 1998: 101. Corydalus camposi Nav as 1935: 368, 369; Contreras-Ramos 1998: 101.
Type locality Panama (Chiriqu ı).
Material examined Antioquia. 1 #, San Luis, Inspeccion de Polic ıa el Vergel, Reserva Ecologica Can~on del R ıo Claro, El Refugio, 425 m, IV.1998, leg. M. Ospina (UNAB). 1 #, Venecia, 14.V.2004, leg. J. Cardona, manual collection (UDEA). 1 #, San Rafael, vereda el Bizcocho, 1100 m, 6.29N, 75W, 26–29.III.2011, leg. C. Florez, mercury light (UDEA- 62008). 1 #,T amesis, vereda La Oculta, finca Kaiser, 5 4505200N, 75 4003800W, 1100 m, 5.VI.2005, leg. E. Alzate, manual collection (UDEA-36786). 1 #, San Jeronimo, VI.1987, leg. A. Arango, manual collection (UDEA). 1 #, Caucasia, hacienda La Candelaria, 8.0302222N, 75.2174389W, 80 m, 15–19.IX.2016, no collector, manual collection (UDEA-94531).
Atlantico. 1 $, Sabanalarga, 8.V.1995, leg. V. Sergio (UNAB). 1 #, Puerto Colombia, 4.III.1920, leg. H. Salazar (MLS-18).
Bol ıvar. 1 #, San Juan de Nepomuceno, 9 5700.0200N, 75 50000W, 297 m, 9.V.2012, leg. A. Ram ırez, manual collection (UNAB). 1 #, San Juan de Nepomuceno, SFF Los Colorados, 9 570N, 75 70W, 180 m, 29.IV.2012, leg. A. Alvarez, entomological net (UNAB).
Caldas. 1 #, Norcasia, Fca. La Germania, I.XI.2014, leg. P. Torres (CAUD).
Caqueta. 1 $, Florencia, IX.1954, no collector (MLS-17).
Casanare. 1 $, Aguazul, 290 m, 5 100N, 72 330W, VI.2012, leg. C. Pimentel, manual collection (UNAB). 1 $, Aguazul, Fca. Camila, 5 160N, 72 280W, 251 m, 19.IX.2012, leg. G. Beltr an, entomological net (UNAB). 1 $, Tauramena, 24.IV.1999, leg. R. Cely (UNAB).
Cundinamarca. 1 #, La Mesa, San Joaqu ın, 10.X.1998, leg. A. Bautista, (UNAB). 1 #, Ricaurte, Vda. La Carrera, 3.IV.1994, leg. N. Bol ıvar (UNAB). 1 #, Mesitas del Colegio, Viot a, fca. Canaima, 4 3104800N, 74 2802600W, 949 m, 20.X2012, leg. J. Navarra, manual collection (UNAB). 1 #, Tocaima, V.28.1967, leg. A. Reyes (UNAB). 1 #, Perto Salgar, Vda. Tres y Medio, Fca. Miraflores, 5 4801700N, 74 6002400W, 177 m, 3.IV.2012, leg. C. Pena,~ manual collection (UNAB). 1 #, Puerto Salgar, Vda. Tres y Medio, Fca. Miraflores, 5 4801700N, 74 6002400W, 177 m, 3.IV.2012, leg. C. Pena,~ 324 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS manual collection (UNAB). 1 #, La Vega, Condomionio Le Porc, 4 110N, 74 270W, 1230 m, 9.V.2010, leg. J. Suescun manual collection (UNAB). 1 $, Anapoima, Vda. San Antonio, 4 330N, 74 320W, 591 m, 26.V.2012, leg. J. Ch avez, manual collection (UNAB). 1 $, Jerusalem, Vda. La Victoria, 4 3304700N, 74 4104300W, 357 m, 4.I.2014, leg. J. Villarraga, manual collection (UNAB). 1 $, Villeta, Vda. Payand e, 5 005300N, 74 2602900W, 845 m, 15.VIII.2012, leg. R. Rojas, manual collection (UNAB). 1 $, Silvania, 23.V.1993, leg. J. Luque (UNAB). 1 $, Guaduas, 5 40N, 74 340W, 2700 m, 18.VIII.2012, leg. D. Munoz,~ entomological net on shrubby vegetation (UNAB). 1 $, Puerto Salgar, Vda. Tres y Medio, 5 290N, 74 390W, 177 m, 4.IV.2012, leg. C. Pena,~ manual collection (UNAB). 1 $, Gachet a, 4 4901500N, 73 3802400W, 1746 m, 11.X.2009, leg. L. Rodriguez, manual collection (UNAB). 1 $, Silvania, Vda. Azaranal, 1500 m, 6.XII.2009, leg. N. Florez, on Coffea arabiga (UNAB). 1 $, Tena, 16.X.1988, leg. F. Jim enez (UNAB). 1 $, Tequendama, Mesitas del Colegio, Fca. El Santuario, 4 340900N, 74 260000W, 1291 m, 15.IV.2010, leg. C. Pinzon, riparian vegetation (UNAB). 1 $, Anapoima, La Paz, 4 50600N, 74 3202200W, 625 m, 20.IX.2004, leg. W. de Leon, manual collection (UNAB). 1 $, Tequendama, Mesitas del Colegio, Fca. El Santuario, 4 340900N, 74 260000W, 1291 m, 11.IV.2010, leg. E. Pinzon, entomological collection (UNAB). 1 $, Anapoima, Vda. Panam a, 700 m, 4 520N, 74 530W, 1.X.2011, leg. L. Soto, manual collection (UNAB). 1 $, Anapoima, Vda. Las Mercedes, 680 m, 1.IV.2010, leg. S. Rodriguez, manual collection (UNAB).
Guain ıa. 1 $, Puerto Inirida, 16.V.1992, leg. S. Bernal (UNAB).
Huila. 1 $, Neiva, Vda. Bajo Bejucal, 2 5905600N, 75 1801600W, 415–440 m, 18–23.VII.2011, leg. S. Castro, entomological net (UNAB).
Meta. 1 #, Villavicencio, Aeropuerto, 467 m, 10.III.2012, leg. R. Pena~ (UNAB). 1 $, Villavicencio, Hotel del Llano, 4 90N, 73 380W, 509 m, 17.III.2012, leg. M. P erez, manual collection (UNAB). 1 $, Villavicencio, Vda. Bueno Aires, Fca. La Esmeralda, 4 70N, 73 390W, 561 m, 2.XI.2014, leg. S. Rodr ıguez, entomological net (UNAB). 1 $, Villavicencio, 4 40N, 73 340W, 947 m, 2.VI.2013, leg. D. Pinzon, manual collection (UNAB). 1 #, Villavicencio, v ıa Acac ıas, Fca. San Andr es, 400 m, 2.V.2010, leg. L. Ariza, manual collection (UNAB). 1 #, San Mart ın, Reserva Las Unamas, 3 540N, 73 110W, 240 m, 11.IV.2011, leg. L. Ojeda, manual collection (UNAB). 1 #, Acac ıas, 5.X.1992, leg. M. Montanez~ (UNAB). 1 #, Granada, 3 340N, 73 430W, 372 m, 23.III.2015, no colector, manual collection (UNAB). 1 $, San Mart ın, 6.IV.1998, leg. L. Piedrahita (UNAB). 1 $, San Mart ın, Reserva Natural el Caduceo, 405 m, 10.XI.2009, leg. J. Delgado, light trap (UNAB). 1 $, San Mart ın, Reserva las Unamas, 3 540N, 73 110W, 240 m, 2.IV.2011, leg. L. Ojeda, light trap (UNAB). 1 $, Cubarral, zona urbana, 25.IV.1992, leg. F. Pel aez (UNAB). 1 $, Guamal, 21.V.2004, leg. R. Lopez (UNAB).
Putumayo. 1 $, La Hormiga, 316 m, 0 250N, 76 540W, 16.XII.2011, leg. Y. Becerra, manual collection (UNAB). 1 #, Puerto Colon, 4.III.1874, leg. H. Salazar (MLS-008).
Santander. 1 $, Contratacion, 6 180N, 73 370W, 1200 m, 7.VI.2003, leg. C. Carranza (UNAB). 1 $, Cabrera, 6 3607.300N, 73 1405800W, 980 m, 17.I.2012, leg. M. Hernandez, AQUATIC INSECTS 325
Figure 6. Corydalus liui sp. n.: (a) habitus of holotype male; (b) head, dorsal view; (c) habitus of paratype female (photos by Francisco Gamarra); (d) mandible of female; (e) head and prothorax of female, dorsal. entomological net (UNAB). 1 $, Socorro, 2.VI.1999, leg. F. Ome (UNAB). 1 $, Barichara, 3.IV.1999, leg. D. Rey (UNAB). 1 #, Florida Blanca, 3.IV.1994, leg. G. Jaimes (UNAB). 1 #, same data but leg. H. Roberto (UNAB).
Tolima. 1 $, Honda, Fca. Santa Ana, 450 m, 5 1202700N, 74 4402800W, 24V.2004, leg. C. Ricaurte, entomological net (UNAB). 1 $, Melgar, 19.XI.1992, leg. I. Rodriguez (UNAB). 1 $, Melgar, Vda. Agua Media, Fca. Santa Luc ıa, 4 100N, 74 340W, 1163 m, 4.III.2012, leg. J. Restrepo, light trap (UNAB). 1 $, Mariquita, Cataratas de Medina, 23–25.X.2001, leg. F. Carranza (UNAB).
Valle del Cauca. 1 $, Sevilla, 25.VII.1990, leg. H. Furque (UNAB).
Diagnosis, adult Forewing length 35.85–59.0 mm in males and 41.0–57.35 mm in females. Head pale brown, with a subtle postocular dark stripe. Male mandible elongated with reduced dentition, and a positive allometric pattern in function of the body size. Male antenna subserrate, typically pale yellow with infuscate apex, sometimes pale to dark brown. Female antenna short, generally dark brown. Forewing pale brown, with distal half, and area surrounding M fork and rarp1 somewhat dark brown pigmented (Figure 2c). Male genitalia with elongated and tubular ectoproct, apical incurvation 326 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 7. Corydalus liui sp. n.: (a–d) male genitalia: (a) ventral view; (b) dorsal view; (c) lateral view; (d) tenth gonocoxites þ gonostyli, ventral; (e, f), female genitalia: (e) ventral view; (f) lateral view; (g, h) male genital primordia: (g) ventral view; (h) dorsal view. Scale bars: 1 mm. well developed; sternite subquadrate with posterolateral lobes moderately developed; gonostylus elongate, subclavate; fused gonocoxites 10 forming a convex plate, antero- medially with a short projection (Figure 3e); gonostyli 10 short, papilliform, slightly divergent (Contreras-Ramos 1998).
Comments Corydalus flavicornis is a common species in Colombia where it has been recorded from Antioquia, Meta, Santander, Bol ıvar, Cundinamarca, Magdalena, Tolima, and AQUATIC INSECTS 327
Figure 8. Corydalus liui sp. n.: (a) male clypeal margin; (b) tenth gonocoxites þ gonostyli, ventral. Drawings by Omar Bernal. Scale bars: 1 mm.
Valle del Cauca departments (Contreras-Ramos 1998; Ardila-Camacho 2014). It is more frequently collected in the Caribbean, Andean and Orinoqu ıa regions of Colombia, while has fewer records from the Pacific region. The altitudinal distribu- tion occurs from 0 to 2700 m, although, the great majority is between 0 and 1200 m, and a few are near 1800 m. The only record over 2700 m is likely erroneous. Adults of this species have been collected throughout the year (Contreras-Ramos 1998). Some aspects of the biology and ecology, as well as a detailed description of the last larval stage were addressed by Azev^edo (2009).
Distribution This species is widely distributed in Central and northern South America, with records from Costa Rica, Colombia, Ecuador, El Salvador, Guatemala, Honduras, Panama, Peru, and Venezuela (Contreras-Ramos 1998).
Corydalus liui sp. n. urn:lsid:zoobank.org:act:E230F184-F695-4D78-97FE-9A9D07EDA87A (Figures 3i, 6, 7, 8, 10, 11)
Type locality Colombia, Santander, Chipat a, 6 0304500N, 73 3802400W, 1700 m.
Type material Holotype. # (reared from the larva), ‘Colombia, Santander, Chipat a, 1700 m, 6 0304500N, 73 3802400W, leg. A. Ardila’ (UNAB), prepupa: 20.I.2016, pupa: 27.I.2016, adult: 16.II.2016. 328 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 9. Corydalus clavijoi Contreras-Ramos, 2002: (a) habitus of holotype male; (b, c) male geni- talia: (b) dorsal view; (c) ventral view. Photos by Marco Gaiani. Paratypes. 1 $, Boyac a, Chitaraque, Vda. Potrero Grande, Fca. La Fr ıa, 1575 m, 6 10N, 73 270W, IV.2010, leg. A. Rojas, light trap (UNAB). 1 $, Cundinamarca, San Antonio del Tequendama, Vda. San Jos e, 4 470700N, 74 1303100W, 1540 m, 18.IV.2015, leg. N. P erez, entomological net (UNAB). 1 #, Cundinamarca, Venecia, La Esmeralda, 2400 m, 4.VIII, leg. J. Mendoza (UNAB). 1 $, Cundinamarca, La Mesa, Vda. San Javier, 1200 m, 11.X.2003, no collector, entomological net (UNAB). 1 $, Cundinamarca, C aqueza, 8.V.1994, leg. F. Martinez (UNAB). 1 #, Cundinamarca, El Colegio, 4 3501400N, 74 2605900W, 5.X.2002, leg. L. Alvarez (CAUD). 1 $ (reared from the larva), Santander, Chipat a, 1700 m, 6 0304500N, 73 3802400W, leg. A. Ardila, larva: 28.XII.2015, pupa: 01.II.2016, adult: 19.II.2016 (UNAB). 1 # (reared from the larva), same data, leg. A. Ardila, prepupa: 20.I.2016, pupa: 25.I.2016, adult: 12.II.2016 (CAUD). 1 $ (reared from the larva), same data leg. A. Ardila, prepupa: 20.I.2016, pupa: 22.II.2016, adult: 8.III.2016 (CAUD). 1 #, Santander, San Jos e de Pare, Region de Ricaurte, Puente Nacional, leg. F. Campos (CAUD). 1 $, Tolima, El L ıbano Km 3, 4 540N, 75 50W, 2000 m, 25.V.2004, leg. C. Valbuena, entomological net (UNAB).
Additional material, adults 1 $, Antioquia, Cocorn a, Vda. El Choco, Fca. ACA, 6 30N, 75 340W, 1300 m, 10.VII.2012, leg. C. Nino,~ entomological net (UNAB). 1 $, Caldas, La Dorada, AQUATIC INSECTS 329
Figure 10. Last instar larva of Corydalus liui sp. n.: (a) habitus, dorsal; (b) habitus, ventral (photos by Francisco Gamarra); (c) head, dorsal; (d) head, ventral; (e) prothorax, dorsal; (f) forefemur, dorsal; (g) mandible; (h) lateral filament of abdominal segment 2; (i) spiracle of the abdominal segment 1; (j) sternellum, lateral; (k) last three abdominal segments, dorsal; (l) macro- and microsetae of the abdominal tergite 5; (m) V-shaped pattern of abdominal sternite 5.
V.1974, leg. A. Lotero (ICN). 1 $, Caldas, Marip ı, 1250 m, V.1940, F. Gallego (MEFLG). 1 #, Santander, San Jos edeSu aita, Vda. La Meseta, Est. Biolog ıa Universidad Nacional, 1500 m, 5.IX.1998, no collector (ICN). 330 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 11. Collecting sites, mature larvae, prepupae, and pupa of Corydalus liui sp. n.: (a) mature larva below of a large stone, in a stream of Chipat a, Santander; (b) mature larva observed in a muddy bottom, in the same locality; (c, d) prepupae in pupal chambers, observed below of a large stone, 1 m away of the water stream, in the type locality (arrowheads); (e, f) type locality of Corydalus liui sp. n.; (g, h) male pupa reared in the laboratory: (g) lateral view; (h) dorsal view. AQUATIC INSECTS 331
Additional material, larvae, prepupae 2 L, Santander, Chipat a, Roperana stream, 1700 m, 6 0304500N, 73 3802400W, 30.VIII.2015, leg. A. Ardila (CAUD). 2 L, same data but 28.V.2015, leg. A. Ardila (CAUD). 5 L, same data but 25.IX.2015, leg. A. Ardila (CAUD). 5 L, same data but 18.XII.2015, leg. A. Ardila (CAUD). 2 L, same data but 28.XII.2015, leg. A. Ardila (CAUD). 3 prepupae, same data but 20.I.2016, leg. A. Ardila (CAUD).
Additional material, pupae See ‘Type material’.
Diagnosis, adult Forewing length 49.13–52.62 mm in males, 48.43–57.65 mm in females. Male man- dible elongated with reduced dentition, flagellum dark brown; male clypeal margin with lateral projections divergent, median projection well developed, medially incised. Forewing with two pale pterostigma-like areas, distal half, radial space, and area sur- rounding M fork with irregular pale areas alternated by pale- to dark brown suffu- sions. Female head dark reddish brown, mandible with a median protrusion contiguous to basal tooth. Male genitalia with ectoproct and gonostylus 9 elongated, tubular slightly subclavate, gonostylus 9 with a small and short preapical claw; sternite 9 modified, with a narrow posteromedian projection, curved dorsad and reaching to middle of gonostyli 10; posterolateral projections well developed, broadened and somewhat subquadrate, sometimes slightly curved mesally. Fused gonocoxites 10 with anterolateral projections elongate and straight, posterolateral sides subquadrate, pos- terior margin straight; gonostyli 10 somewhat elongated, blunt, sclerotized, divergent.
Measurements Male. Mandible length, 10.11–13.69 mm; head width (at the level of postocular flange), 6.34–7.09 mm; head length (clypeal margin–occipital suture), 5.86–6.27 mm; prothorax length, 4.60–5.36 mm; pterothorax length, 5.81–7.45 mm; forewing length, 49.13–52.62 mm; hindwing length, 44.17–48.40 mm; antenna length, 29.99–35.41 mm.
Female. Mandible length, 4.85–5.17 mm; head width (at the level of postocular flange), 6.30–7.50 mm; head length (clypeal margin-occipital suture), 5.97–6.94 mm; prothorax length, 5.06–5.57 mm; pterothorax length, 6.53–8.47 mm; forewing length, 48.43–57.65 mm; hindwing length, 45.70–51.81 mm; antenna length, 20.77–21.55 mm.
Description of adult Head. Subquadrate, pale brown, without muscle scars; male mandible elongated, den- tition not completely reduced (Figure 6b), with small preapical teeth, basal protrusion slightly developed; maxilla brown, maxillary palpi with four palpomeres, the last with a partial subdivision, dark brown; metal plates brown, submental projections blunt, not surpassing anterior margin of mental plates, labial palpi 3–articulate, the last art- icle with an ellipsoid sensory area, dark brown. Clypeal margin medially moderately incised, lateral projections developed, divergent (Figure 8a); gena brown with anterior margin pale brown. Ocelar region very dark brown, compound eyes metallic green. 332 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Postocular region infuscated; postocular flange and spine, moderately developed. Filiform antenna, scape and pedicel brown, 60–63 flagellomeres dark brown, covered with short spiniform setae.
Thorax. Prothorax subcyllindrical, slightly longer than wide, pronotum subrectangu- lar, pale brown (Figure 6a), covered with scattered fine setae, basisternum pale brown. Meso- and metanotum brown, covered with fine golden setae. Pteropleura ligth brown, with some dark brown spots on posterior region of metapleura, entire surface densely covered by long golden setae.
Legs. Brown, except apex of tibiae, and tarsi, which are dark brown; all segments cov- ered by short brown setae, tarsal claws brown.
Wings. Forewing elongate, ellipsoid, membrane pale brown, semi-translucent; costal space pale- to dark brown, with 36–40 crossveins, dark brown with a central pale area, apical region with two pale pterostigma-like areas; subcostal space pale- to dark brown, with single basal crossvein. Area between RA and Cu with diffuse and irregu- lar pale areas surrounded by pale- to dark brown suffusions; area adjacent to M fork with dark brown maculation (Figure 6a); RP with seven or eight branches, and 15 or 16 crossveins, dark brown; longitudinal veins pale brown. MA with single fork, MP unforked; area between M and Cu with four or five crossveins; CuA with four branches. Hindwing elongate, subelliptical, membrane pale brown, semi-translucent (Figure 6a); costal field 24–29 crossveins, pterostigmal area pale brown; longitudinal veins pale brown; area between RA and MA pale brown; RP with eight branches, 14–16 crossveins; MA with two branches, MP unforked; area between M and CuA with four or five crossveins; CuA with four branches.
Abdomen. Abdominal segments 1–8 dark brown (Figure 6a), with fine short golden setae, tergite 9 pale brown; sternites pale brown, with fine short golden setae.
Male genitalia. Tergite 9 medially subdivided, laterally with two bands of fine and short setae, posterolaterally slightly projected (Figure 7b). Sternite 9 subtrapezoidal, posteromedially with an elongate and narrow projection, nearly as long as tergite length, curved and extended until mid-length of gonostyli 10, apex blunt (Figure 7a); posterolateral lobes developed, subquadrate, densely covered by long and fine setae (Figure 7a); membrane between sternite 9 and gonocoxites 10 bilobed, somewhat sclerotized. Ectoproct elongate, tubular, slightly longer than gonostylus 9, basal 1/3 slightly broadened, apex subclavate, entire surface covered by fine long setae (Figure 7b). Gonostylus 9 elongate, tubular, curved dorsad, distal half slightly broadened, with a small preapical claw, entire surface densely covered by long and fine setae (Figures 7a–c). Gonocoxites 10 not completely fused, with a medial subdivision, anterolateral projections long, broad, straight or medially slightly curved, with blunt apex, posterior margin subquadrate (Figures 7d, 8b). Gonostyli 10 papilliform, scler- otized, slightly divergent, ventrally projected, somewhat elongate and covered by fine and short setae (Figures 7d, 8b). AQUATIC INSECTS 333
Female. Head dark reddish brown (Figure 6e); mandible basally yellow, first preapical tooth as long as mid-length of apical tooth, second and third preapical teeth short, similar in size, basal protrusion forming an obtuse angle with the inner margin of the mandible (Figure 6d). Clypeal margin piceous, medially emarginated, incised with developed posterolateral projections. Gena with yellow margins. Postocular region with a subtle dark stripe; postocular flange and spine developed. Filiform antenna with 54–61 flagellomeres, black, covered by short spiniform setae. Pronotum dark brown (Figure 6e); meso- and metanotum pale brown laterally with two dark brown spots, scutellum brown, covered by fine and long golden setae; pteropleura brown with dark brown spots, covered by long and fine golden setae. Legs brown, except fore coxa, fore femur apex, and mid- and hindtibia dark reddish brown, tarsal claws dark reddish brown, all segments covered by abundant brown setae. Forewing colour pattern similar to male but darker (Figure 6c); costal field with 42–45 crossveins; Rs with 8 branches, 16–20 crossveins. Hindwing costal space with 27–31 crossveins; Rs with seven or eight branches, 14–18 crossveins. Abdominal terga dark brown, stern- ites pale brown (Figure 6c). Gonocoxite 8 sclerotized, posterior margin straight (Figure 7f), in ventral view anterior margin concave, posteromedially strongly sclerot- ized, slightly incised (Figure 7e); tergite 9 posteroventrally projected in lateral view, covered by fine, long setae (Figure 7f); ectoproct in lateral view posterodorsally with a rounded lobe; gonocoxite 9 short, covered by long and fine setae (Figure 7f).
Comments In a recent work on Colombian Corydalus, Ardila-Camacho (2014) reported the first records of Corydalus clavijoi Contreras-Ramos, 2002 for the country, a species previ- ously known solely from the Merida Andes of Venezuela. The species was reported from a series of specimens from central Colombia, in the eastern slope of the western cordillera. After the examination of several additional specimens from entomological museums and specimens reared in the laboratory, it was determined that the sup- posed specimens of C. clavijoi actually belong to a new species. The wing and anten- nal colour patterns, plus male mandible morphology of both sexes of the new species are very similar to those of C. clavijoi (Figure 9a), however the morphology of the male ectoproct and gonostylus 9 are different, being wider and slightly apically sub- clavate in the new species. The preapical claw of the gonostylus 9 is smaller and short in C. liui (Figures 7a, b), while it is elongate and prominent in C. clavijoi (Figures 9b, c). Furthermore, anterolateral projections of the gonocoxites 10 are straight, postero- lateral corners are broad and subquadrate and the posterior margin is straight. In constrast, C. clavijoi has anterolateral projections of gonocoxites 10 laterally curved, and the posterior part of these sclerites is narrower and rounded; gonostyli 10 of the new species are blunt and slightly elongate, well sclerotized and divergent, and in C. clavijoi they are shorter and subparallel. With this finding, the number of species of the C. nubilus species group increases from four to five representatives, although formal phylogenetic analyses to establish the relationships between these species are required. As mentioned above, C. liui is apparently the vicariant sister species of C. clavijoi, rather than C. tesselatus Stitz, 1914 as it was previously hypothesized by Contreras-Ramos (2011). Another 334 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS distinctive member of this group is C. muzoensis Ardila-Camacho, 2014, also discov- ered from a neighboring mountainous region in the central Colombian Andes. That species is probably the sister taxon of the Venezuelan species C. tesselatus, and, if so, the idea of an origin by means of geographical isolation of those species pairs appears very likely.
Etymology This species is named after Xingyue Liu of China Agricultural University of Beijing, who has extensively contributed to the knowledge of Megaloptera. The name is a noun in the genitive case.
Diagnosis, larva Body length 42.91–64.46 mm, mandible narrow, basal half yellow to pale brown, basal tooth closer to apex than base, third preapical tooth not completely fused to basal tooth; submental projections blunt, slightly surpassing the anterior margin of the mental plates; area between ocular region and frontal sclerite with a conspicuous sub- trapezoidal yellow spot, posterior area to frontal sclerite with two subtriangular yellow spots. Sternellum short and blunt with a few short and thick setae. Abdominal terga densely covered by scale-shaped decumbent microsetae, and scattered subclavate stri- ated black macrosetae, medial region with numerous long and tapered pale brown setae; lateral filaments long and thickened, similar in shape and size on the different segments, except that of the first abdominal segment which has nearly half the length of the remainder filaments; all the filaments dorsally with three thick rows of hair- like setae.
Description of last larval instar Head. Labrum pentagonal, dark amber at margins, yellow at centre, anterior edge with two pairs of long setae; clypeus subtrapezoidal, pale amber, except lateral mar- gins amber; postclypeus dark brown with two lateral yellow spots that are extended towards gena (Figures 10, 11). Mandible somewhat narrow, dark reddish brown, with centre and base yellow to pale amber, apical tooth long, pointed, first and second pre- apical teeth slightly shorter than apical tooth; third preapical tooth reduced, rounded; basal tooth closer to apex than mandible base, not fused to the third preapical tooth (Figure 10g). Gena dark brown with a semicircular yellow spot, sometimes completely dark brown. Frontal sclerite dark brown with three yellow muscle scars. Antenna short, with five antennomeres, pale- to dark brown. Area immediately posterior to frontal sclerite with two semi-triangular yellow spots (Figure 10c). Area between frontal sclerite and stemmata with a conspicuous subtrapezoidal yellow spot (Figure 10c). Adjacent area to occipital and coronal sutures punctate, very dark brown, with yellow to pale brown muscle scars. Ocular region comprising six stemmata, arranged as a pentagon, with a single central stemma; area adjacent to corneal lenses very dark brown. Postocular area dark brown with yellow muscle scars, lateroventrally with a longitudinal dark brown stripe; postocular keel very dark brown, with a row of short and thick pale amber setae, posterior area to postocular keel with pale yellow muscle scars, and a longitudinal dark brown stripe; area between gular plate and post ocular AQUATIC INSECTS 335 area yellow, with some dark brown infuscations, entire surface with some scattered and short brown setae. Occiput alternating very dark brown and pale yellow. Cardo semicircular, bicoloured, dark amber on inner half, pale amber on outer half; stipes with outer half amber, remainder of surface yellow, inner margin with a row of short and thick setae; lacinia and maxillary palpus amber. Labium mainly yellow, with mar- gins amber; ligula and basal part of palpi covered by short and thick brown setae, labial palpi with three palpomeres amber in colour; prementum pale brown, mental plates mainly pale brown with dark amber margins (Figure 10d), gular plate pale brown, bordered with dark amber, and with a medial dark brown stripe extended from the middle of it until anterior margin where is laterally projected; submental projections somewhat elongate, slightly surpassing anterior margin of mental plates, with blunt apex (Figure 10d).
Thorax. Pronotum subquadrate, glabrous, very dark brown with pale brown muscle scars, and two yellow stripes on lateral margins (Figure 10e). Meso- and metanotum subrectangular, very dark brown, with pale brown muscle scars (Figure 10e); pleural membrane yellow with some brown infuscations, entire surface covered scale-shaped decumbent microsetae, and some scattered subclavate black macrosetae. Prosternum yellow to pale brown, with lateral dark brown spots, sternellum short and blunt, cov- ered with short and thick brown setae (Figure 10j). Meso- and metasternum yellow with brown infuscations, entire surface covered by scale-shaped microsetae.
Legs. Coxae yellow to pale brown, amber at margins, entire surface with short and thick brown setae. Trochanter yellow, amber at base, covered by short and thick brown setae, basally with a tuft of brown setae. Femora, ventrally yellow with numer- ous short and thick brown setae; dorsally dark brown with some longitudinal pale brown stripes, mid-line with a row of long and fine brown setae, remainder of surface covered by short and thick brown setae (Figure 10f). Tibiae colour and setation simi- lar to femora, except foretibia that lacks of dorsal row of long setae, meso- and meta- tibia with a row of shorter setae. Fore leg with femur and tibia bearing prominent setae on inner surface. Tarsi blackish brown on dorsal surface, brown on ventral sur- face, apex with fine and long setae, tarsal claws amber.
Abdomen. Somewhat flattened, tergites dark brown, spiracles anterolaterally situated on each segment (Figure 10a), beside of each spiracle and close to lateral filament bases a tuft of long and fine, pale brown setae present; medial region of abdominal tergites with numerous fine and long yellowish setae, the remainder of surface, including intertergal membranes densely covered by scale-shaped, clear, decumbent microsetae, and scattered, subclavate, dark macrosetae (Figure 10l). Spiracles reni- form, with a yellow concentric stripe, dark brown at centre and periphery (Figure 10i). Lateral filaments long and thickened, tapering, similar in shape and size in the different segments, except those of the first abdominal segment that reach approxi- mately half the length of the remainder filaments (Fig 10a), all of them dorsally dark brown with three thick rows of long and fine setae (Figure 10h); ventrally pale brown (Figure 10b), and covered by subclavate dark macrosetae, and fine and short setae on 336 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 12. Corydalus sophiae sp. n.: (a) habitus of holotype male (photos by Francisco Gamarra); (b–d, g) male genital structures: (b) male genitalia, ventral; (c) same, dorsal; (d) same, lateral; (e) head and thorax, dorsal; (g) tenth gonocoxites þ gonostyli, ventral; (f) male mandible. the remainder of surface. Sternites yellow (Figure 10b), laterally with a pair of long tapering setae, medial region with a V-shaped glabrous region (Figure 10m), remain- der of surface covered by scale-shaped clear microsetae; ventral tracheal gills yellow. Sternites of abdominal segments 9 and 10 with a thick brush of brown setae. Anal AQUATIC INSECTS 337
Figure 13. Corydalus sophiae sp. n.: (a) male clypeal margin, dorsal; (b) tenth gonocoxi- tes þ gonostyli. Drawings by Omar Bernal. Scale bars: 1 mm. prolegs dark brown, with a layer of scale-shaped microsetae, and some subclavate dark macrosetae; cerci with a row of fine and long setae (Figure 10k); claws amber.
Description of pupa Body length 59.40 mm, head subquadrate, yellow, postocular flange and spine slightly developed, mandibles mainly yellow, apically dark brown, antenna yellow, coiled; compound eyes weakly developed, dark brown. Prothorax subquadrate (Figure 11h), yellow with fine pale brown setae; pterothorax yellow to pale brown, wing pads yel- low to pale brown, according to developmental phase; legs yellow. Abdomen, soft, ventrally curved, tergites very dark brown (Figures 11g, h), densely covered by fine golden setae; spiracles brown; sternites pale yellow or cream; pleural membrane bicol- oured with cream and dark brown. Male genital primordia with tergite 9 subtrapezoi- dal in dorsal view, densely setose (Figure 7g); ectoproct short, lobe shaped, ventromedially curved with abundant pilosity; gonostylus 9 reduced; fused gonocoxi- tes þ gonostyli 10 plate-shaped, arched, medially subdivided, gonostylus 10 short and lobe-shaped (Figure 7h).
Bionomics The larval description of this species was made from a series of specimens collected in a small montane stream located over 1700 m a.s.l. in the Chipat a Municipality (Santander Department). The specimens were found while collecting C. armatus lar- vae from the same stream, but, each species is apparently occupying different eco- logical microhabitats. For example, C. armatus was found in a wide range of microhabitats, including stones, logs, decomposing plant material, and submerged roots, either in open sunny areas or in shaded zones with abundant riparian vegeta- tion. In that sense, C. armatus is likely a generalist species. By contrast, C. liui larvae were always found in places with very specific environmental conditions, generally in 338 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS shaded and cold zones, with abundant riparian vegetation, platforms of sedimentary rock, shallow water flux, availability of submerged plant roots, and decomposing leaves (Figures 11e, f). These larvae were generally located beneath flat fragments of sedimentary rock or associated with submerged plant material. Several pre–pupae were found beneath stones in very dry soil beside the water (Figures 11c–f). Interestingly, exuviae of C. armatus pupae were also collected in neighboring habitats.
Corydalus sophiae sp. n. urn:lsid:zoobank.org:act:359A7EEF-0668-408E-B387-E9AC93B3A1D7 (Figures 12, 13)
Type locality Colombia, Antioquia, San Roque, Vereda el Espinal, CPA, San Jos e del Nus, 870 m.
Type material Holotype. #, ‘Antioquia, San Roque, Vereda el Espinal, CPA, San Jos e del Nus, 870 m, 6.XI.2014, leg. J. Ospina’ (MEFLG). Holotype condition: good, pinned, wings extended, lacking antennae, terminalia dissected and cleared, stored in a glass vial filled with glycerin and pinned beneath the specimen.
Diagnosis Forewing length 59.26 mm; head narrow, dark reddish brown, postocular ridge and spine moderately developed. Male mandible short, with well-developed dentition, inner margin without protuberances. Forewing strongly patterned, wing membrane dark brown with numerous irregular pale areas, surrounded by a blackish brown suf- fusion; area surrounding first 1ra–rp and M fork dark brown. Male genitalia with ectoproct short, densely setose, basal half subcylindrical, progressively narrowing towards the apex, slightly flattened; gonostylus 9 elongate, slightly longer than ecto- proct, distinctively subclavate, densely setose; sternite 9 subquadrate with posterolat- eral lobes slightly developed; fused gonocoxites 10 forming a moderately sclerotized plate, with anterolateral projections short, with a rounded lobe opposite to the gono- styli 10; gonostyli 10 papilliform, semi-membranous, short, widely separated, slightly convergent, not surpassing the posterior margin of gonocoxites 10.
Measurements Mandible length, 4.50 mm; head length (occiput-clypeal margin), 7.00 mm; head width (at the level of postocular ridge), 6.00 mm; pronotum length, 4.77 mm; pterothorax length, 8.00 mm; forewing length, 59.26 mm; hindwing length 52.66 mm.
Description Head. Small, dark reddish brown (Figure 12e); male mandible short, with well-devel- oped dentition, without protuberances, dark reddish brown, except by an orange sul- cus extended from between basal and second preapical teeth to outer margin, and base yellow; basal tooth situated at the level of mandible midlength, first and second preapical teeth subequal, apical tooth long and pointed (Figure 12f). Maxilla dark AQUATIC INSECTS 339 reddish brown, maxillary palpi with four palpomeres, the last palpomere partially sub- divided. Labium dark reddish brown, mental plates small, dark reddish brown on posterior half, brown on anterior half. Submentum dark reddish brown, submental projections long, pointed, surpassing the anterior margin of mental plates. Labrum subrectangular, anterolaterally rounded, with a slight concavity, predominantly dark reddish brown. Clypeal margin with lateral projections moderately developed, medial projection wide with medial incision (Figure 13a). Gena with anterolateral margin orange. Scape and pedicel pale brown. Dorsal ocelli with orange corneal lenses. Postocular ridge and spine moderately developed.
Thorax. Prothorax elongate, pronotum subrectangular, dark reddish brown, without distinct muscle scars, laterally covered by fine brown setae, medial region glabrous (Figure 12e). Mesonotum nearly as long as wide, dark brown, scutellum with poster- ior margin pale brown, entire surface covered with numerous fine, and long black setae. Metanotum mainly brown, anteriorly with a mixture of tan and brown, entire surface with abundant long and fine black setae. Pteropleura brown, sclerite margins tan, entire surface covered with long and fine black setae.
Legs. Fore coxa dark reddish brown, mid- and hind coxae brown, all of them covered by long and fine black setae. Trochanter and femur in all legs pale brown. Fore tibia dark brown on inner surface, outer surface brown, mid- and hind tibia pale brown. Foretarsi tan, brown on mid- and hind leg; tibial spurs short, tarsal claws amber. All segments covered with short brown setae.
Wings. Forewing elongate, subelliptical, wing membrane semi-translucent; costal space broad, brown, with 25 or 26 crossveins, of which 4 or 5 are forked, all of them dark brown with surrounding membrane dark brown; cells on the medial region with semicircular blackish spots, on some cells together semicircular, fenestrate, white spots; pterostigmal area slightly marked, composed by 14 or 16 crossveins. Subcostal space pale brown with dark brown infuscations, a single basal crossvein with a dark brown macula. Area between RA and posterior wing margin strongly patterned with numerous irregular pale areas, surrounded by a dark brown suffusion (Figure 12a); membrane surrounding 1ra–rp and M fork dark brown. RP with five or six branches, 11–14 crossveins. Longitudinal veins pale brown with blackish tips; crossveins black- ish brown with surrounding membrane dark brown; basal portion of CuP and CuA branches with dark brown spots. Area between M and CuA with four crossveins; CuA with four branches. Hindwing elongate, pale brown, semi-translucent; costal space narrow with 40 crossveins, the first four or five pale brown, the remainder dark brown, some cells with diffuse dark brown spots. C and Sc veins mainly brown, other longitudinal veins pale brown with blackish tips, except anal veins; basal portion of CuA branches dark brown. Crossveins dark brown with diffuse dark brown spots on membrane. Radial cells with small irregular pale areas bordered with dark brown suf- fusion (Figure 12a). RP with five or six branches, 11 crossveins, cells with diffuse pale areas. MA with two branches, MP unforked. Area between M and CuA with four or five crossveins; CuA with four branches. 340 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Abdomen. Dark brown, covered with fine, short, amber setae.
Male genitalia. Tergite 9 subtrapezoidal, medially subdivided in a pair of hemiter- gites, posterolaterally slightly produced, each hemitergite covered by abundant pilosity (Figure 12c). Membrane between sternites 8 and 9 medially forming a small pocket. Pregenital sacs scarcely developed. Ectoproct elongate, tubular, slightly longer than tergite 9, basal half subcylindrical with lateral margins subparallel, distal half, some- what flattened, progressively narrowed towards the apex, entire surface densely setose (Figure 12c); sternite 9 subquadrate, anterior margin concave, posterolateral lobes slightly developed, entire surface densely covered by long and fine setae (Figure 12b); gonostylus 9 elongate, densely setose, subclavate, slightly longer than the ectoproct, in lateral view slightly curved dorsad (Figure 12d); membrane between sternite 9 and gonocoxites 10, developed, bilobed, moderately sclerotized; fused gonocoxites 10 forming a sclerotized plate, with anterolateral projections short, somewhat curved, blunt, followed by a rounded projection nearly opposite to the gonostyli 10 (Figures 12g, 13b); gonostyli 10 papilliform, short, nearly as long as wide, not surpassing pos- terior margin of the gonocoxites 10, convergent and covered by short setae (Figures 12g, 13b). Area posterior to gonocoxites 10 covered by abundant long setae (Figure 12g).
Comments Corydalus sophiae belongs to a group of atypical species distributed in the western region of northern South America. The group includes C. colombianus and C. ecua- dorianus Banks, 1948 from the rainforest of the Colombian Pacific coast and the mountainous area of the Ecuadorian Andes, respectively. These species are distin- guished by their strongly patterned wings, showing numerous irregular pale areas sur- rounded by dark brown pigmentation. In C. colombianus, the head is moderately patterned, with some clear muscle scars in a dark background; the male mandible is elongate with discrete dentition, and a conspicuous protrusion at mid-length on inner margin is present. On the other hand, C. ecuadorianus has the head with a contrast- ing colour pattern, and the muscle scars are quite conspicuous; the male mandible is short with well-developed dentition, and there is a slight basal protrusion, close to the basal tooth. By contrast, the head of C. sophiae does not have a contrasting colour pattern, and is completely dark reddish brown; the male mandible is short and simi- lar to that of C. ecuadorianus, but a basal protrusion is lacking in C. sophiae. The wing colour pattern of the new species is noticeably darker and more elaborate than that of C. colombianus. Moreover, the morphology of the male genital structures is more similar to that of C. ecuadorianus, suggesting these two species are sister taxa. Although both species have similarly shaped ectoprocts, the morphology of the gono- stylus 9 is quite distinct, being tubular and digitiform in C. ecuadorianus and subcla- vate in C. sophiae. The gonocoxites þ gonostyli of both species are quite similar as well, but the gonostyli 10 of C. sophiae are remarkably separated and convergent. In addition, the new species has the anterolateral projections of the gonocoxites 10 very distinct from that of C. ecuadorianus, particularly because the presence of a rounded lobe apposite to the gonostyli 10. AQUATIC INSECTS 341
This species group diverged early in the phylogeny of the genus, together with C. cephalotes, C. arpi, and the C. nubilus groups. These taxa are morphologically heterogeneous, and may represent a remnant of diverse archaic clades; or, they may represent species that evolved in isolation (Contreras-Ramos 1998). Corydalus sophiae probably represents the sister taxon of C. ecuadorianus, and both must have diverged across the geological history of the mountainous ranges of the Andes.
Etymology This species is named in special honor of Lauren Sophia, daughter of the first author. The name is a noun in the genitive case.
Corydalus muzoensis Ardila-Camacho, 2014 (Figures 2d, 3h) Corydalus muzoensis Ardila-Camacho 2014: 112.
Type locality Colombia (Boyac a).
Diagnosis, adult Forewing length 45.0 mm in male. Head small, dark reddish brown, postocular spine short. Antennae filiform, thin, dark reddish brown, with spinous setae. Male mandible elongated, with discrete dentition. Forewing semi-translucent, costal space, medially infuscate, two pale pterostigma-like areas present; area between RA and CuA with brown suffusions; radial cells with a few semicircular fenestrate white spots; M fork with surrounding membrane dark brown (Figure 2c). Male genitalia with elongate and slightly flattened ectoproct, mesally curved inwards; gonostylus 9 short, dorsome- dially projected, basal half wider than distal half, apex constricted forming a small bulb, abruptly bent inwards at an angle of 90 and with an apical claw (Figure 3h); sternite 9 concave, posteromedially with a long acuminate projection, dorsally curved, reaching to the level of the midlength of gonostyli 10, posterolateral lobes well devel- oped, slightly mesally curved (Figure 3h); gonocoxites 10 forming a plate, medially not completely fused, and with anterolateral projections long, sinuous and pointed; gonostyli 10 elongate, digitiform, slightly curved ventrad, strongly sclerotized and sharply pointed (Figure 3h).
Comments Species known only from the type locality, the Muzo Municipality, Boyac a Department. The only known altitudinal records is over 850 m a.s.l. The specimen was collected in the vicinity of tributaries of the Minero River during July. The female and immature stages are unknown.
Distribution Known from central Colombia, on the western slopes of the eastern cordillera, in the Boyac a Department. 342 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Corydalus nubilus Erichson, 1848 (Figures 2e, 3g) Corydalis nubila Erichson 1848: 586. Corydalus titschacki Nav as 1928b: 60, 61, 62; Geijkes 1984: 79.
Type locality Guyana (no further locality data).
Material examined 1 #, Vaup es, Mitu, v ıaCano~ Sangre, 26–27.IV.2014, 200–300 m, entomological net near water stream, 10:00 h, E. Realpe (CAUD).
Diagnosis Forewing length 47.7–61.0 mm in males and 54.0–63.0 mm in females. Head and pro- notum pale to moderately dark brown, with conspicuous lateral dark stripe. Male mandible elongated with reduced dentition with a positive allometric pattern in func- tion of the body size, basal protrusion conspicuous. Antenna filiform, thin, black. Forewing pale yellowish brown, semi-translucent with subtle dark brown suffusions between RA and CuA (Figure 2e). Male genitalia with elongate ectoproct, slender with convex inner surface at base, apex subclavate; gonostylus 9 elongated, as long as half-length of ectoproct, distal half claviform; sternite 9 semi-triangular with a short posteromedian projection, posterolateral lobes absent (Figure 3g); fused gonocoxites 10 forming a moderately convex plate, with anterolateral projections short, bluntly rounded; gonostyli 10 elongated, subrectangular, dorsal- and apically sclerotized (Contreras-Ramos 1998). The last larval instar with a body length of 52.50–61.70 mm; head and pronotum from pale brown to reddish brown. The mandible with the most basal tooth nearer to apex than base; submental projections short with acute apex, not surpassing or reaching the level of the anterior margin of the mental plates. The sternellum is triangular, short and blunt, without an apical curvature, and covered with short, thick setae. The abdominal terga bear tubular and elongate macrosetae that are brown to black in colour (Azev^edo and Hamada 2006, 2014).
Comments This species is distributed in lowland forests of the Amazon basin. Adults have been collected nearly throughout the year except in February, June, July, and December, and records from August and November are most common (Contreras-Ramos 1998). In Colombia, it has been reported from Vaup es Department (Ardila-Camacho 2014). The last larval instar and bionomic data were described by Azev^edo and Hamada (2006), while some aspects of its ecology were studied by Azev^edo (2009).
Distribution Brazil, Colombia, French Guiana, and Venezuela. AQUATIC INSECTS 343
Corydalus peruvianus Davis, 1903 (Figures 2f, 3f, 14) Corydalis peruviana Davis 1903: 480. Corydalus primitivus fera Nav as 1927: 30; Contreras-Ramos 1998: 152.
Type locality Guatemala (no further locality data).
Material examined Antioquia. 1 #, Cocorn a, Vda. El Choco, Fca. ACA, 6 30N, 75 340W, 1300 m, 1.VIII.2012, leg. C. Nino,~ manual collection (UNAB). 1 #, Antioquia, V. Vergara (UNAB); 1#, Ciudad Bolivar, Farallones, Vda. La Mina, Quebrada Buenarenton, 5.80368 N, 76.0691 W, 2000 m, 29.V.2011, leg. J. Uribe, manual collection on stream margin (UDEA). 1 #, Cocorn a, Finca La Golosa, 7.XII.2007, leg. A. Montoya (UDEA). 1 #, Cocorna, 15.XI.1997, leg. M. Duque, attracted to light (UDEA). 1 #, T amesis, VII.2002, leg. J. V elez (UDEA). 1 #, vereda La Oculta, finca Kaiser, 5 4505100N, 75 4003700W, 1100 m, 5.VI.2005, 20:14h, leg. E. Alzate (UDEA36783). 1 #, Quimbaya, vereda Quemon Km. 7, parque Panaca, 1200 m, Leg. C. Pablo, manual collection (UDEA-40493). 1 #, Maceo, Vereda San Pedro, 6 4605500N, 74 7906000W, 800 m, 1.IV.2011, leg. N. Urquijo, manual collection at night (UDEA-62012).
Boyaca. 1 #, Soat a, Vda. La Casta sector Villa Guas abara, 6 190N, 72 400W, 1470 m, 28.X.2013, leg. J. Sandoval (UNAB).
Caldas. 1 #, La Dorada, 26.XI.1994, leg. H. Bejarano (UNAB). 1 #, Saman a, casco urbano, 5 29044.3000N, 75 0051.7800W, 1004 m, 2.X.2012, leg. L. P erez, manual collec- tion (UNAB).
Cauca. 1 #, Tierradentro, 3.V.1994, leg. C. Acuna~ (UNAB).
Casanare. 1 #, Aguazul, 16.X.1995, leg. T. corredor (UNAB).
Cundinamarca. 1 #, Guayabetal, 11.XI.1995, leg. M. Moya (UNAB). 1 #, Quipile ori- ente, Fca. Campo Alegre, 1500 m, 31.X.2009, leg. F. Zubieta, manual collection (UNAB). 1 #, Sasaima, Santa Ana, 31.I.1998, leg. V. Bernal, K. Turriago (UNAB). 1 #, Tibacuy, La Vuelta, Fca. La Esperanza, 3.X.2009, 1800 m, leg. R. Simbaqueva, man- ual collection (UNAB). 1 #, Fusagasug a, camino a la aguadita, 4 2004900N, 74 2105300W, 1731 m, 15.IV.2005, leg. N. Orjuela (UNAB). 1 #, Fusagasug a, XI.2009, leg. L. Betancourt, manual collection (UNAB). 1 #, Girardot, 12.X.1992, leg. A. Escobar (UNAB). 1 #, Anapoima, Vda. Panama, 4 520N, 74 530W, 700 m, 14.II.2011, leg. C. Soto, entomological net (UNAB). 1 $, Villeta, Miradores del Payand e, vda. El Naranjo, 18.I.2013, leg. M. Parada, (CAUD). 344 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
Figure 14. Corydalus peruvianus Davis, 1903, females observed in field: (a) females and egg masses on a large stone, inside a river of Santa Mar ıa, Boyac a; (b) aggregation of females laying egg masses in the same locality. Photos by Diego Gomez.
Huila. 1 #, Pitalito, Vda. Aguadas, Parque Agroecologico Yamboro, 1 530N, 76 50W, 1290 m, 25.IX.2014, leg. J. Navas, manual collection (UNAB).
Meta. 1 #, San Mart ın, Reserva Caduceo, 3 3901900N, 73 3405100W, 400 m, 1–3.XI.2011, leg. N. Salcedo, entomological net (UNAB). 1 #, Granada, casco urbano, 332 m, 3 3204000N, 73 4202300Wm 9.IV.2012, leg. Y. Mar ın, manual collection (UNAB). 1 #, Villavicencio, Hotel Ciudad, 4 90N, 73 370W, 467 m, 18.V.2013, leg. A. Forero, attracted to light (UNAB). AQUATIC INSECTS 345
Santander. 1 #, Barichara, 31.III.1999, O. Higuera, R. Quevedo (UNAB). 1 #, Barichara, 10.IV.1999, leg. C. Moreno (UNAB). 1 #, Barichara, 18.IV.1999, leg. J. Afanador (UNAB). 1 #, Barichara, 6.VI.1999, leg. J. Arbel aez (UNAB). 1 #, Contratacion, 6 1705900N, 73 2804600, 1690 m, 28.V.2004, leg. J. Garzon, manual collec- tion on Citrus sp. (UNAB). 1 #, Guavat a, 6.X.1995, no collector (UNAB). 1 #, San Jos e de Pare, 17.IV.1993, leg. H. Rodriguez (UNAB). 2 #,V elez, 19.XI.2013, on a white wall during daytime, leg. A. Ardila (CAUD).
Tolima. 1 #, Flandes, 6.I.1995, V. Wilches (UNAB). 1 #, Mariquita, V.1999, leg. E. Gordillo (UNAB).
Valle del Cauca. 1 #, Yotoco, Reserva Natural Yotoco Universidad Nacional, 3 5205.500N, 76 2301,7500’W, 1200 m, 7.X.2012, leg. J. Guerrero, manual collection (UNAB). 1 #, Yotoco, R.N. Yotoco Universidad Nacional, 3 5205.500N, 76 2301,7500W, 1200 m, 7.X.2012, leg. J. Guerrero, manual collection (UNAB). 2 #, Calima, 2.VI.1969, no collector (MLS-001, 002).
Diagnosis Forewing length 44.9–77.75 mm in males and 50.0–70.9 mm in females. The head generally dark greenish brown, without muscle scars. The male mandible elongated with reduced dentition and has a positive allometric pattern as a function of the body size. Male antennae are thickened, slightly sub-serrate, pale yellow to greenish yellow, and the apex is infuscate (Figure 2f). The forewing is dark greenish to greyish brown, more or less uniformly coloured, and with small semicircular fenestrate white spots (Figure 2f). The male genitalia have elongated and tubular ectoproct, with a devel- oped apical incurvation; gonostylus 9 elongate, subclavate; sternite 9 subquadrate, posterolateral lobes moderately developed; fused gonocoxites 10 forming a somewhat convex plate, with anterolateral projections long and narrow, nearly straight; gonostyli 10 nearly two times as long as wide, with about half of gonostylus surpassing the pos- terior margin of gonocoxites 10 (Figure 3f); pregenital sacs absent (Contreras- Ramos 1998).
Comments In Colombia this species was previously reported from Antioquia, Cundinamarca, Meta, Putumayo, Valle del Cauca, Caldas, Casanare, Cauca, Guajira, Boyac a, Santander, and Tolima, making it one of the most common dobsonfly species in the country (Contreras-Ramos 1998; Ardila-Camacho 2014). Altitudinal distribution ranges from 40 to 2700 m a.s.l., but the vast majority of records are reported below of 1000 m. Adults have been collected throughout the year.
Bionomics An aggregation of females of C. peruvianus was sighted in the Municipality of Santa Mar ıa, Boyac a laying their eggs on large stones along a montane stream (Figure 14a). The females laid their eggs in consecutive rows, and the latter were covered by a whitish substance that later hardened (Figure 14b). Once the females finished the 346 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS oviposition, they remained in the same place, possibly protecting the egg masses until they had hardened. These egg masses have a similar appearance to bird droppings. Such whitish substance has a protective function against desiccation and predators, although some insects in the families Trichogrammatidae (Hymenoptera), Chloropidae (Diptera), and Cantharidae (Coleoptera) are known to be egg predators or parasitoids of Corydalus (Azev^edo 2009).
Distribution Argentina, Bolivia, Colombia, Costa Rica, Ecuador, Guatemala, Mexico, Panama, Peru, and Venezuela (Contreras-Ramos 1998).
Key to Colombian species of Megaloptera (After Penny and Flint 1982; Contreras-Ramos 1998, 2002; Ardila-Camacho 2014; Liu et al. 2015c, modified)
1. Forewing length 7.0–14.5 mm; ocelli absent; fourth tarsomere bilobed...... (Sialidae, Ilyobius)2 – Forewing length 24.0–85.0 mm; ocelli present; fourth tarsomere simple, unmodi- fied ...... (Corydalinae) 3 2. Head blackish brown with dark orange on medial part of vertex ...... I. flavicollis – Head mainly orange with a pair of blackish brown spots surrounding the com- pound eyes, extended towards dorsal and posterior cephalic regions (Figure 1a)...... I. curvata 3. Body colouration pattern pale yellow with black spots; forewing with last RP branch forked (Figures 1b, c) ...... Chloronia 4 – Body colouration pattern from pale brown to dark reddish brown; forewing with last RP branch generally unforked (Figures 1d–f) ...... Corydalus 7 4. Male sternite 9 with posterior and lateral margins produced into two elongate lobes ...... 5 – Male sternite 9 with two well-developed lateral lobes, posteromedial lobes not developed...... C. bogotana 5. Male gonostylus 9 incurvate, with widened basolateral angle; gonostyli 10 mamilliform...... C. gloriosoi – Male gonostylus 9 slightly basally narrowed; gonostyli 10 semi–triangular, strongly pointed ...... 6 6. Male ectoproct bearing an apical group of long and thick setae; gonostylus 9 with a preapical tuft of long and thick setae (Figure 3a)...... C. marthae – Male ectoproct with an apical group of short and thick setae; posteroventral margin of gonostylus 9 with a longitudinal row of long setae...... C. mirifica 7. Forewing dark brown, strongly patterned with numerous irregular pale areas surrounded by a dark brown suffusion ...... 8 – Forewing pale to dark brown, generally with a colour pattern more or less uni- form, or with a discrete to moderately strong pigmentation pattern, composed AQUATIC INSECTS 347
of pale to dark spots and irregularly shaped pale areas, or with small semicircu- lar fenestrate white spots ...... 9 8. Head colour pattern uniform, dark reddish brown; male mandible short, with well-developed dentition (Figure 12f); gonostylus 9 elongate, subclavate, without apical claw (Figure 12b); gonostyli 10 papilliform and slightly convergent (Figure 13b) ...... C. sophiae sp. n. – Head with a contrasting colour pattern, with slightly defined muscle scars; male mandible elongate with discrete dentition and conspicuous basal protrusion (Figure 2b); gonostylus 9 elongate, subcylindrical, narrow and with a small apical claw; gonostyli 10 broad, subquadrate and divergent...... C. colombianus 9. Sternite 9 modified, posteromedially sclerotized with a developed projection, posterolateral projections generally well developed ...... 10 – Sternite 9 typically subquadrate ...... 13 10. Head and pronotum with a lateral dark stripe (Figure 2e); sternite 9 posterome- dially slightly produced, posterolateral lobes not developed (Figure 3g); gonosty- lus 9 subclavate with a basodorsal sclerotized lobe...... C. nubilus – Head and pronotum with uniform colour pattern; sternite 9 with a prominent posteromedial projection, posterolateral lobes well developed; gonostylus 9 with an apical or preapical claw ...... 11 11. Sternite 9 with a thin posteromedial projection, posterolateral lobes broad, sub- quadrate; gonostylus 9 elongate, tubular with a small preapical claw (Figures 7a–c)...... C. liui sp. n. – Sternite 9 with a prominent posteromedial projection, acuminate or thumb-like, posterolateral lobes directed mesally; gonostylus 9 with an apical claw ...... 12 12. Sternite 9 with a thumb–like posteromedian projection; gonostyli 10 short, sub- quadrate, sclerotized...... C. tesselatus – Sternite with a prominent acuminate posteromedian projection (Figure 3h); gon- ostyli 10 elongate, digitiform, strongly sclerotized, distinctively curved ventrad with pointed apex ...... C. muzoensis 13. Head pale brown with a subtle lateral dark brown stripe; forewing pale brown with irregular semi-translucent pale areas (Figure 1d); ectoproct and gonostylus 9, subequal in shape and length, elongate and narrow (Figure 3b)...... C. affinis – Head colour pattern uniform, varying from pale brown to dark reddish brown; forewing with numerous small semicircular fenestrate white spots, sometimes with dark maculations on adjacent area to M fork; ectoproct tubular generally with an apical incurvation, gonostylus 9 subclavate to subglobose ...... 14 14. Forewing with a contrasting colour pattern, with distal half, portions of costal space, radial cells, and adjacent area to M fork being dark reddish brown (Figure 1f) ...... C. batesii – Forewing colour pattern slight- to moderately marked, pale- to dark brown ... 15 15. Forewing with distal half and adjacent area to rarp1 cell and M fork moderately darkened (Figure 2c); fused gonocoxites 10 with an anteromedian projection (Figure 3e)...... C. flavicornis – Forewing pale- to dark brown, often nearly uniformly pigmented (Figures 1e, 2a); fused gonocoxites 10 without anteromedian projection (Figures 3c, f) ...... 16 348 A. ARDILA-CAMACHO AND A. CONTRERAS-RAMOS
16. Head with a subtle postocular vitta; antenna very dark brown to black, shiny (Figure 2a); forewing pale- to dark golden brown (Figure 2a); gonostylus 9 sub- globose, posterior area to gonocoxites 10 with long hair–like setae..... C. clauseni – Head colour pattern uniform; male antenna pale- to dark greenish yellow; fore- wing pale brown to greenish brown or dark greyish brown; gonostylus 9 elong- ate and subclavate; posterior area to gonocoxites 10 with short setae ...... 17 17. Male antenna (including scape and pedicel) pale to dark brown, apically infus- cated (Figure 1e); gonostyli 10 nearly as long as wide, with less than half length surpassing the posterior margin of gonocoxites 10 (Figure 3c); pregenital sacs well developed and conspicuous (Figure 3c) ...... C. armatus – Male antenna (including scape and pedicel) yellow to greenish yellow, with apical 1/3 infuscated (Figure 2f); gonostyli 10 nearly two times as long as wide, with nearly a half-length surpassing posterior margin of gonocoxites 10 (Figure 3f); pregenital sacs apparently absent (Figure 3f) ...... C. peruvianus
Key to last larval instar to families, genera and some species of Colombian Megaloptera (After Azev^edo and Hamada 2014, modified)
1. Abdominal segments with eight pairs of lateral filaments, ventral tracheal gills on segments 1–7 present, last segment with a pair of false prolegs equipped with claws ...... (Corydalidae, Corydalinae) 2 – Abdominal segments with seven pairs of lateral filaments, tracheal gills absent, last segment with a long tapered filament ...... (Sialidae) Ilyobius 2. Body length 20.0–50.0 mm; head and pronotum without muscle scars, pronotum with two pairs of dark spots, sometimes fused forming two lateral stripes, or completely dark reddish brown ...... Chloronia – Body length 40.0–80.0 mm; head and pronotum with a conspicuous pattern of muscle attachment scars (Figures 4a, 10a) ...... Corydalus 3 3. Mandible with basal tooth closer to the base than apex ...... 7 – Mandible with basal tooth closer to apex than base ...... 4 4. Submental projections short, not surpassing the anterior margin of mental plates (Figure 4d); area between the stemmata and frontal sclerite pale- to dark brown, with diffuse pale brown spot (Figure 4c) ...... 5 – Submental projections elongated, slightly surpassing the anterior margin of men- tal plates (Figure 10d); area between stemmata and frontal sclerite with a con- spicuous subtrapezoidal yellow spot (Figure 10c) ...... C. liui sp. n. 5. Abdominal tergites with elongate macrosetae ...... C. nubilus – Abdominal tergites with subglobose, claviform macrosetae ...... 6 6. Sternellum elongate, curved, projected towards anterior part; abdominal tergites with dark, claviform macrosetae ...... C. affinis – Sternellum short, blunt, ventrally projected (Figure 4i); abdominal tergites with dark, subglobose macrosetae (Figure 4k) ...... C. armatus 7. Area between stemmata and frontal sclerite with a diffuse yellow spot; mandible with pointed basal tooth ...... C. flavicornis AQUATIC INSECTS 349
– Area between stemmata and frontal sclerite with a wide yellow vitta, extended towards anterior part of frontal sclerite; mandible with basal tooth broad and bluntly rounded ...... C. batesii
Acknowledgements We offer our sincere thanks to Francisco Serna of Facultad de Agronom ıa, Universidad Nacional de Colombia, sede Bogot a, Erika Valentina Vergara of Facultad de Agronom ıa, Universidad Nacional de Colombia, sede Bogot a, and Coleccion Taxonomica Nacional de Insectos ‘Luis Mar ıa Murillo’ Corpoica, Mosquera, Cundinamarca, Emilio Realpe from Universidad de los Andes, Bogot a, Alexander Garc ıa from Universidad Distrital, Bogot a, Fernando Sarmiento of the Universidad de la Salle, Bogot a, Nancy Carrejo, Museo Entomologico of the Universidad del Valle, Cali, Marta Wolff from the Universidad de Antioquia, Medell ın, John Albeiro Quiroz of the Museo Entomologico “Francisco Luis Gallego”, Universidad Nacional de Colombia, sede Medell ın, Fernando Fernandez and Carlos Sarmiento of the Instituto Nacional de Ciencias Naturales, Universidad Nacional de Colombia, sede Bogot a for allowing us to revise the specimens deposited under their care. Thanks to Robin Thomson, University of Minnesota, St. Paul, Marco Gaiani, Universidad Central de Venezuela, Caracas, and Xingyue Liu, China Agricultural University, Beijing for their kindness to share photographs on type specimens of Megaloptera from Colombia and Venezuela. The first author is in much indebted to Diego Gomez, Universidad de los Andes for share photo- graphs and data of Megaloptera taken during his field trips in Colombia. Sincere acknowledg- ment to Francisco Gamarra, Universidad de los Andes for contributing the majority of the photographs used herein, and Omar Bernal, Universidad Pedagogica Nacional, Bogot a for the drawings of the genitalia used in this study.
Disclosure statement No potential conflict of interest was reported by the authors.
ORCID Adrian Ardila-Camacho https://orcid.org/0000-0002-3750-867 Atilano Contreras-Ramos http://orcid.org/0000-0001-8044-1348
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