New, additional and historical records of Leptophlebiidae (Insecta, Ephemeroptera) from Colombia

In the Neotropical region, especially South America, Leptophlebiidae achieves its higher diversity, with 45 genera and almost 250 species. In Colombia, the diversity about this family is underestimated, with only 22 species reported so far. In order to contribute to the current knowledge of the Leptophlebiidae in Colombia we examined the adults of this family deposited at the Colección Entomológica del Programa de Biología, Universidad de Caldas. In addition to the historical records herein presented, nine species are registered for the first time from the country (Farrodes savagei, F. tepui, F. xingu, Hagenulopsis minuta, Terpides contamanensis, T. echinovaris, T. ornatodermis, Tikuna atramentum, and Ulmeritoides flavopedes). Besides that, additional records are presented for 12 species of the following genera: Askola, Farrodes, Simothraulopsis, Tikuna, Ulmeritoides, Hydrosmilodon, Terpides, Choroterpes, Paramaka, and Hagenulopsis. With these additional and new records, we update the list of Leptophlebiidae from Colombia to 33 species and 14 genera. A R T I C L E I N F O Article history: Received 28 June 2020 Accepted 10 August 2020 Available online 7 September 2020 Associate Editor: Douglas Zeppelini


Introduction
Leptophlebiidae is one of the most diversified groups of mayflies (Ephemeroptera), both taxonomically and ecologically, with more than 640 species and 140 genera distributed worldwide (Sartori and Brittain, Salles et al., 2020b). These numbers, especially for Leptophlebiidae and Baetidae, are far from the actual number. In South American countries where the mayfly components have been studied more often, such as Brazil and Argentina, the number of species of this families surpasses one hundred and many species remain to be described.
Based on material collected by L.G. Dias team in the last years and deposited at the Entomological Collection of the Biology program of the Caldas University -CEBUC, our aim is to increase the knowledge of Leptophlebiidae from Colombia by presenting new and additional records of the family from the country.

Occurrence maps
We made the maps of occurrence in South America with the software ESRI® ArcMap 10.0, using a digital model of the elevation with countries delimitation in the Neotropic. Records from the literature were obtained from the following publications: Needham and Murphy (1924); Traver and Edmunds (1967); Savage and Peters (1983); Zúñiga et al. (1997); Mosquera et al. (2001); Domínguez and Zúñiga (2003)

Images
We took pictures of specimens deposited in alcohol using LEICA M205 A and LEICA M205 C stereo microscopes with LEICA MC170 HD digital cameras. Pictures of living specimens in the field were taken with a Nikon d800 camera, 105mm macro lens, a SB-5000 Nikon flash, and a small acrylic aquarium. Post-processing of pictures were made in Adobe Lightroom® and the final plates were prepared in Adobe Illustrator® and Adobe Photoshop®.

Deposition
We deposited the material in the Colección Entomológica del Programa de Biología, Universidad de Caldas (CEBUC), Manizales, Colombia, and at the Museu de Entomologia, Universidade Federal de Viçosa (UFVB), Viçosa, Brazil. In the case of Terpidinae, all specimens are temporarily deposited at the UFVB.
A complete list of the species of Leptophlebiidae reported from Colombia is presented below. For species reported for the first time from the country, the specific name is succeeded by an asterisk (*). Additional records from Colombia: none.

Discussion
As expected, Hagenulinae is the most species-rich subfamily in Colombia, currently represented by 10 genera and 25 species. Among them, Farrodes (five species), Miroculis (four) and Thraulodes (four) stand out as the most diverse, closely followed by Atopophlebia and Hagenulopsis (three species each). In the case of Thraulodes, however, it should be noted, as previously stated, that additional species will be treated elsewhere. Terpidinae is represented by its three genera and seven species, while Choroterpinae is represented by a single new species of Choroterpes. Unidentified species of this genus have been previously reported from Colombia based on nymphs (Peters et al., 2005;Gutiérrez and Reinoso-Flórez, 2010;García et al., 2013), and for the first time adults were found. These are the only records of the world-widely distributed Choroterpinae in South America. The absence of Atalophlebiinae in Colombia, at least among the previous and new records, was somewhat expected. Members of this subfamily include cold-adapted mayflies Monjardim et al., 2020) that, in South America, are mostly restricted to Patagonia and some marginal areas along the southern portion of South American Transition Zone  (the only exception is the genus Massartella Lestage, found in Brazil and Venezuela). It is very unlikely, therefore, that even additional collections in the Paramo, an area poorly represented at the CEBUC, would lead to the discovery of some Atalophlebiinae in the country.
Of the 33 species of Leptophlebiidae treated in this work, 14 are registered only to Colombia while 19 are present in other countries, such as Brazil (eight), Peru (five), Costa Rica (five), and Panama (four). Given the current distribution of these species, the fauna of Leptophlebiidae in Colombia seems to be a mosaic of elements from distinct biogeographic dominions (see Morrone, 2014). Some endemic, such as Atopophlebia fortunensis, Farrodes caribbianus, and Hydrosmilodon primanus (from the Pacific Dominion) or Askola emmerichi, Ulmeritoides flavopedes, and Hagenulopsis minuta (from the Boreal Brazilian Dominion). While others are present in more than one biogeographic dominion. For example, Fittkaulus amazonicus (northwestern portion of the South Brazilian dominion and Boreal Brazilian), and Terpides contamanensis and Terpides echinovaris (northwestern portion of the South Brazilian Dominion and Pacific).
The new and additional records presented in this work contribute significantly to our understanding of the knowledge of the order Ephemeroptera from Colombia. Furthermore, this work highlights the richness of the different biogeographical regions of Colombia and confirms that diversity in this country is underestimated. Our results also demonstrate the importance of biological collections as a depository of biodiversity and suggest that the knowledge of Ephemeroptera in Colombia could expand with the increase in the sampling effort, since many regions of the country have not been studied.

Acknowledgments
We thank the Universidad de Caldas for providing funds to LGD to conduct the sampling of Ephemeroptera in several projects. We also thank Dr. Tito Bacca (Universidad del Tolima), Milton Montaño, MSc Camilo Llano, MSc Ana Meza and students of the Laboratorio de Colecciones Biológicas of Universidad de Caldas for assistance with sampling specimens deposited in CEBUC collection. We thank ISAGEN for facilitating the sampling of macroinvertebrates in the east of the department of Caldas. We also thank Dr. Og DeSouza (Universidade Federal de Viçosa) for facilitating the use of optic equipment. This study was financed in part by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) to JFM (process # 162813/2018-0) and to FFS (processes # 305847/2015-5 and 309666/2019-8).

Conflicts of interest
The authors declare no conflict of interest.

Author contribution statement
LGD conceived the research, conducted the field work and take some imagens. FFS identified the species and elaborated the discussion. ICH obtained some imagens and produced the final plates, and included the world distribution of the lineages. JFM wrote the document and elaborated the maps. All authors critically reviewed the manuscript and approved the final version of this paper.