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The Ephemeroptera Cretaceous Research 127 (2021) 104923 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes The Ephemeroptera (Hexapoda, Insecta) from the Lower Cretaceous Crato Formation (NE Brazil): a new genus and species, and reassessment of Costalimella zucchii Zamboni, 2001 and Cratogenites corradiniae Martins-Neto, 1996 * Natalia C.A. Brandao~ a, b, , Jonathas S. Bittencourt b, Adolfo R. Calor c, Marcio Mendes d, Max C. Langer e a Programa de Pos-graduaç ao~ em Zoologia, Instituto de Ci^encias Biologicas, Universidade Federal de Minas Gerais, Av. Antonio^ Carlos 6627, 31270-901, Belo Horizonte MG, Brazil b Laboratorio de Paleontologia e Macroevoluçao,~ CPMTC, Departamento de Geologia, Instituto de Geoci^encias, Universidade Federal de Minas Gerais, Av. Antonio^ Carlos 6627, 31270-901, Belo Horizonte MG, Brazil c Laboratorio de Entomologia Aquatica (LEAq), Instituto de Biologia, Universidade Federal da Bahia, Av. Barao~ de Jeremoabo, 147, Campus Ondina, 40170- 115, Salvador BA, Brazil d Laboratorio de Paleontologia, Centro de Ci^encias, Universidade Federal do Ceara, campus do Pici, 60455-760, Fortaleza CE, Brazil e Departamento de Biologia, Faculdade de Filosofia, Ci^encias e Letras de Ribeirao~ Preto, Universidade de Sao~ Paulo, Av. Bandeirantes 3900, 14040-901, Ribeirao~ Preto SP, Brazil article info abstract Article history: A new genus and species of Ephemeroptera, Astraeoptera cretacica gen. et sp. nov., is described from the Received 3 December 2020 Lower Cretaceous limestone of the Crato Formation (Brazil). The new taxon has the following diagnostic Received in revised form characters: veins MP2 e CuA straight at their bases, MA branching in the apical half of wing length, CuA not 5 May 2021 forked, cubital field with longitudinal veins sub-parallel to CuA. It was compared with other adult mayflies Accepted in revised form 29 May 2021 worldwide, both fossil and extant, and could not be assigned to any known ephemeropteran family. In Available online xxx addition, holotypes of other two mayfly species were reanalyzed, resulting in taxonomical changes. A new description of the Costalimella zucchii Zamboni, 2001 holotype is provided, indicating its placement within Keywords: Aquatic insects Hexagenitidae, rather than Siphlonuridae, as previously proposed. We also reanalysed the holotype of Fossil Cratogenites corradiniae Martins-Neto, 1996, providing new and more accurate information about the Mayfly species and confirming its junior synonymy to Protoligoneuria limai Demoulin, 1955. Our results indicate Paleontology that the diversity of Crato Formation mayflies, both at species and family level, is greater than previously Gondwana thought, confirming the predominance of hexagenitids among Cretaceous ephemeropterans. Taxonomy © 2021 Elsevier Ltd. All rights reserved. 1. Introduction approximately 3500 extant species described so far (Sartori and Brittain, 2015). The phylogenetic position of the group is contro- Ephemeroptera is a widespread order of amphibiotic insects versial and has been discussed on the basis of three hypothesis: (i) (Dominguez et al., 2006; Da-Silva and Salles, 2012), popularly as closely related to Odonata (Palaeoptera hypothesis; Martynov, known as mayflies. Their temporal record spans minimally from 1925; Crampton, 1924; Haas and Kukalova-Peck, 2001; Kukalova- Permian (Grimaldi and Engel 2005; Staniczek, 2007) to recent, with Peck, 2008; Regier et al., 2010; Blanke et al., 2013; Thomas et al., 2013; Song et al., 2019), (ii) as closely related to Neoptera (Chias- tomyaria hypothesis; Carle, 1982a, 1982b; Kjer, 2004; Yoshizawa * Corresponding author. Programa de Pos-graduaç ao~ em Zoologia, Instituto de and Johnson, 2005; Misof et al., 2007; Simon et al., 2009), and Ciencias^ Biologicas, Universidade Federal de Minas Gerais, Av. Antonio^ Carlos 6627, (iii) sister group of clade Odonata þ Neoptera (Metapterygota hy- 31270-901, Belo Horizonte MG, Brazil. pothesis; Staniczek, 2000; Wheeler et al., 2001; Ogden and ~ E-mail addresses: [email protected] (N.C.A. Brandao), [email protected] Whiting, 2003; Grimaldi and Engel, 2005; Willkommen and (J.S. Bittencourt), [email protected] (A.R. Calor), [email protected] (M. Mendes), Hornschemeyer,€ 2007; Zhang et al., 2008). Despite this long [email protected] (M.C. Langer). https://doi.org/10.1016/j.cretres.2021.104923 0195-6671/© 2021 Elsevier Ltd. All rights reserved. N.C.A. Brandao,~ J.S. Bittencourt, A.R. Calor et al. Cretaceous Research 127 (2021) 104923 history of disagreement, there is a general consensus that mayflies The Crato Formation paleoenvironment has been variously constitute an early branch of winged insects (Grimaldi and Engel, considered as a lake or lagoon, but recent data based on multiproxy 2005; Dominguez et al., 2006; Staniczek, 2007; Sartori and analyses suggest that this Lagerstatte€ is better interpreted as a Brittain, 2015). seasonal, semi-arid, shallow lacustrine wetland (Ribeiro et al., The paleodiversity of mayflies is particularly noticeable during 2021). The age of the Crato Formation has been considered as late the Cretaceous, with 50 genera and 69 species described from all Aptian in most papers dealing with the biostratigraphy of the continents, except Antarctica, a distribution that is coincident with Araripe Basin (Ponte & Appi, 1990; Arai et al., 2001; Coimbra et al., that of extant Ephemeroptera (Sartori and Brittain, 2015). Fossil 2002; Heimhofer & Hochuli, 2010; Coimbra & Freire, 2021). A mayflies are especially diverse in the Early Cretaceous, partly due to recent work suggested a latest Barremianeearliest Aptian deposi- their abundance and exquisite preservation in the Aptian strata of tion for the black shales of the Ipubi Formation (overlying the Crato the Crato Formation (Araripe Basin), northeast Brazil. Fifteen Formation), based on ReeOs absolute ages (Lúcio et al., 2020). mayfly species have been described from this lithostratigraphic Based on those isotopic data, the authors proposed a unit, including representatives of the families Australiphemeridae Barremianeearly Aptian age for the Crato Formation. Yet, this (Australiphemera revelata McCafferty, 1990 and Microphemera neo- proposition has been criticized due to its inconsistency with the tropica McCafferty, 1990), Baetiscidae (Protobaetisca bechlyi bioestratigraphic data (Coimbra, 2021) and is open to scrutiny. Staniczek, 2007), Ephemeridae (Cratonympha microcelata Martins- Neto and Caldas, 1990), Euthyplociidae (Pristiplocia rupestris 3. Material and methods McCafferty, 1990), Hexagenitidae (Protoligoneuria limai Demoulin, 1955, Cratohexagenites longicercus Staniczek, 2007 and Cratohex- The specimens studied in this paper are housed in the fossil agenites minor Staniczek, 2007), Oligoneuriidae (the recently collections of the Laboratorio de Paleontologia de Ribeirao~ Preto described Incogemina nubila Storari et al., 2020, Colocrus magnum (LPRP/USP), Universidade de Sao~ Paulo, in Ribeirao~ Preto, Brazil, Staniczek, 2007 and Colocrus indivicum McCafferty, 1990, recently and Laboratorio de Paleontologia, Centro de Ciencias,^ Universidade reclassified as an hexagenitid by Storari et al., 2020), Poly- Federal do Ceara (LP/UFC), in Fortaleza, Brazil. mitarcyidae (Caririnympha mandibulata Martins-Neto and Caldas, Photographs were taken with a Leica DFC 295 camera, coupled 1990), Potamanthidae (Olindinella gracilis Martins-Neto and to Leica M205C stereomicroscope. Image processing was made Caldas, 1990), and possibly Siphlonuridae (Costalimella nordestina with Leica Application suite LAS v.3.8 and line drawings using a Martins-Neto, 1996 and Costalimella zucchii Zamboni, 2001). In fact, lightbox and later digitized and edited using Adobe Illustrator CC several previously proposed species not listed above have been 2017. The descriptive terminology, including wing venation, follows synonymized in the last years (see review of Staniczek, 2007). This Kluge (2004). is the case of the putative hexagenitid Cratogenites corradiniae This work has been registered in the Official Register of Martins-Neto, 1996, synonymized with Protoligoneuria limai by Zoological Nomenclature (ZooBank) with LSID urn:lsid:zoo- Staniczek (2007). bank.org:pub:77D723D8-55B1-4F55-94EB-DF746F59EC9F. The above-mentioned record represents ca. 7.5% of the 121 insect families of the Crato Formation (Moura-Júnior et al., 2018). Also 4. Taxonomy according to those authors, more than 100 species have been described in the last decade, which demonstrates the enduring Class Insecta Linnaeus, 1758 potential of the Cretaceous strata of the Araripe Basin for taxonomic Order Ephemeroptera Hyatt and Arms, 1891 studies of insects. Despite the significant, already known diversity, Suborder Euplectoptera Tillyard, 1932 some of which requires new investigations, there are hundreds of Family Incertae sedis mayfly specimens from the Crato Formation deposited in research Genus Astraeoptera gen. nov. institutions in Brazil and abroad still awaiting study. Those have the (LSID: urn:lsid:zoobank.org:act:24C6C337-C976-4D63-9336- potential to reveal an even greater taxonomic and morphological BA14F125D0BA). diversity. In order to expand the available data about the ephemer- Type species: Astraeoptera cretacica sp. nov., by monotypy. opteran diversity in the Early Cretaceous, we describe a new species and genus that could not be assigned to any of the known mayfly Etymology. The generic name derives from Latin Astræus (Greek: families and redescribe the holotypes of Cratogenites
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