Phylogeny and classification of Leptophlebiidae (Ephemeroptera) with an emphasis on Neotropical fauna

Abstract

Mayflies from the family Leptophlebiidae are cosmopolitan and highly diverse morphologically; they are also the largest family in numbers of genera and the second in number of species in the order Ephemeroptera. In spite of their broad diversity and the efforts employed to understand the evolution of this group, the internal classification of Leptophlebiidae remains controversial at all levels. More recently, important changes have been incorporated into the systematics of the family, increasing the number of subfamilies (from two to six) and recognizing several tribes. We present a phylogeny of the family based on 153 taxa (53 genera) and two molecular markers, representing 1655 bp, and verify the taxonomic status of the subfamilies, tribes and complexes. Based on these results, the number of subfamilies has been increased from six to eight and one new tribes and two new subtribes have been added. In addition, new ranks are proposed and the concept of Atalophlebiinae revised, including genera with distributions in the Australasian and Neotropical regions.

Gondwana breakup under the ephemeral look

“…The topology of Atalophlebiinae herein proposed contains four main lineages: Garinjuga from Australia, Massartella from South America, and two others with genera from South America and Australia region (Figures 1, 2, and S1). Garinjuga samples and new sequences addition (Table ) helped to establish the relationship among main lineages of the group, which in previous research were unclear (see Monjardim et al, 2020). Jappa and Austrophlebioides Campbell & Suter, 1988, belong to the clade supported by node 8 (Figures 1 and 2) diverging from the results of Monjardim et al, (2020).…”

“…We also used 59 sequences available on GenBank to complement our dataset (Table ). Nine outgroup genera were selected based on a recent phylogeny of the Ephemeroptera (Monjardim et al, 2020; Ogden et al, 2019; O'Donnell & Jockusch, 2008) and the availability of fossils for calibration. Our data matrix comprised 19 genera, representing approximately 55% of the generic composition of Atalophlebiinae.…”

“…Atalophlebiinae (Ephemeroptera, Leptophlebiidae) are a cool‐adapted mayfly subfamily with amphinotic distribution, present in temperate and mountainous areas of South America and Australia (Monjardim et al, 2020; O'Donnell & Jockusch, 2008; Pescador & Peters, 1980; Savage, 1987). Previously, researchers also considered taxa from tropical areas of South America, Africa, and Madagascar as part of this group, suggesting an ancient Gondwana origin of this subfamily (Kluge, 2009; Pescador & Peters, 1980; Savage, 1987).…”

“…Previously, researchers also considered taxa from tropical areas of South America, Africa, and Madagascar as part of this group, suggesting an ancient Gondwana origin of this subfamily (Kluge, 2009; Pescador & Peters, 1980; Savage, 1987). However, recent studies indicate a monophyletic lineage of only amphinotic taxa (Monjardim et al, 2020; O'Donnell & Jockusch, 2008). Ephemeroptera is one of the oldest insect lineages (Misof et al, 2014), and its amphinotic distribution pattern is recurrent in other taxa of the group, such as Ameletopsidae, Coloburiscidae, Nesameletidae, and Oniscigastridae (see Edmunds, 1972; Sartori & Brittain, 2015).…”

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

“…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 (Santos et al, 2018;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 (Molineri et al, 2020) (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.…”