Spathiurus dorsalis Davis, 1887, from the Upper Cretaceous of Haqel, Lebanon, and the evolutionary history of Ionoscopiformes Tamara El Hossny a, b, c, * , Sibelle Maksoud c, d , Dany Azar c, d , Pierre Abi Saad e , Lionel Cavin a a Natural History Museum of Geneva, Department of Geology and Paleontology, CP 6434, Geneva 6,1211, Switzerland b University of Geneva, Department of Earth Sciences, Rue des Maraîchers 13, Geneva, CH-1205, Switzerland c Lebanese University, Faculty of Science II, Fanar, Natural Sciences Department, Fanar e Matn, PO Box 26110217, Lebanon d Chinese Academy of Sciences, Nanjing Institute of Geology and Palaeontology, Nanjing, 210008, People’s Republic of China e Memory of Time, Citadel Area, Byblos, 26111, Lebanon article info Article history: Received 19 March 2020 Received in revised form 27 July 2020 Accepted in revised form 14 August 2020 Available online 21 August 2020 Keywords: Haqel Cenomanian Ionoscopiformes Ionoscopidae Halecomorphi Spathiurus Amphilaphurus abstract New and well-preserved fossil fish material from the Upper Cretaceous (Cenomanian) of Haqel, Lebanon, is assigned to Spathiurus dorsalis Davis, 1887. Spathiurus dorsalis and Amphilaphurus major are found to be synonymous. A complete re-description of this taxon is provided and delivers new recognized anatomical information, mainly cranial features. Spathiurus dorsalis displays diagnostic characters of ionoscopids and consequently is designated as a new member of this family. A phylogenetic analysis was performed to place this taxon within the Halecomorphi. The studied taxon was recovered as the sister genera of Ionoscopus petrarojae, both derived compared to ‘Ionoscopus’ cyprinoides, suggesting that the latter should be placed in a distinct genus. The validated placement of Spathiurus within the Ion- oscopidae extends the stratigraphic range of this family from the Albian to the Cenomanian and becomes the first record of an ionoscopiform from the Upper Cretaceous and from the Middle East. © 2020 Elsevier Ltd. All rights reserved. 1. Introduction Halecomorphs, a clade of neopterygian fishes gathered with the ginglymodians among the holosteans, are represented today by a single living species, the bowfin Amia calva Linnaeus, 1766. In contrast with their poor diversity of the present day, their fossil record is highly diverse (e.g., Parasemionotiformes, Ophiopsidae, Ionoscopidae, Oshuniidae, Caturidae, Sinamiidae, Amiidae, etc.) (Grande and Bemis, 1998). Within the Halecomorphi, the order Ionoscopiformes includes three families: Ophiopsidae Bartram, 1975, Oshuniidae Grande and Bemis, 1998 and Ionoscopidae Lehman, 1966 (Grande and Bemis, 1998). This order was later divided into two families Ionoscopidae and Ophiopsidae by Alvarado-Ortega and Espinosa-Arrubarrena (2008). Ebert (2018) grouped the Ionoscopidae within the Amiiformes thus rejecting the order Ionoscopiformes, and the new order Ophiopsiformes Ebert, 2018, was erected including the new family Ophiopsiellidae Ebert, 2018. He discarded the Ophiopsidae that appears to be non- monophyletic. Taxa worth mentioning are included in the Ophiopsiellidae by Ebert (2018): Agoultichthys chattertoni Murray and Wilson, 2009; and Petalopteryx syriacus Pictet, 1850. Murray and Wilson (2009) actually described A. chattertoni as a macro- semiid based on the presence of a long dorsal fin, short jaw and absence of fulcra in the pectoral fins. They also found similarities between Placidichthys and the macrosemiids, as already pointed out by Brito (2000), based on the presence of two dorsal fins, which is a shared unique feature among the non-teleostean actino- pterygians. Petalopteryx syriacus was related by Pictet (1850) to the living flying gurnard (Dactylopterus) and assigned to the Macro- semiidae by Hay (1903), whereas Woodward (1942) considered it as an amiid rather than a macrosemiid. L opez-Arbarello and Sferco (2018) found in their recent study dealing with the Neopterygii relationships that the order Ion- oscopiformes is not monophyletic. Machado (2016) also doubted the monophyly of this order and found the ionoscopids to be more closely related to the Amiiformes than to the ophiopsids, as Ebert * Corresponding author. E-mail address: Tamara.El-Hossny@ville-ge.ch (T. El Hossny). Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes https://doi.org/10.1016/j.cretres.2020.104619 0195-6671/© 2020 Elsevier Ltd. All rights reserved. Cretaceous Research 116 (2020) 104619