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The First Fossil Owl (Aves, Strigiformes) from the Paleogene of Africa diversity Article The First Fossil Owl (Aves, Strigiformes) From the Paleogene of Africa N. Adam Smith 1,* , Thomas A. Stidham 2,3,4 and Jonathan S. Mitchell 5 1 Campbell Geology Museum, Clemson University, Clemson, SC 29634, USA 2 Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China; [email protected] 3 CAS—Center for Excellence in Life and Paleoenvironment, Beijing 100044, China 4 University of Chinese Academy of Sciences, Beijing 100049, China 5 West Virginia University Institute of Technology, Beckley, WV 25801, USA; [email protected] * Correspondence: [email protected] Received: 10 April 2020; Accepted: 21 April 2020; Published: 23 April 2020 Abstract: The relatively extensive fossil record of owls (Aves, Strigiformes) in North America and Europe stands in stark contrast to the paucity of fossil strigiformes from Africa. The first occurrence of a fossil owl from the Paleogene of Africa extends the fossil record of this clade on that continent by as much as 25 million years, and confirms the presence of large-sized owls in Oligocene continental faunas. The new fossil is tentatively referred to the Selenornithinae, a clade of large owls previously restricted to Europe. This new fossil owl was likely similar in size to the extant Eagle Owls of the genus Bubo, and suggests that the niche of large, volant, terrestrial avian predator, although relatively rare throughout avian evolutionary history, may be an ecological role that was more common among extinct owls than previously recognized. Keywords: Egypt; Fayum Depression; Jebel Qatrani Formation; Oligocene; Selenornithinae 1. Introduction Owls (Strigiformes) are a morphologically distinctive, geographically widespread, and relatively diverse clade with a fossil record stretching back more than fifty-five million years [1–3]. Owls are terrestrial predators that occupy higher trophic categories and comprise >200 extant species around the globe, being absent today only in Antarctica and a few isolated islands [1]. The fossil record of owls is quite rich overall, including approximately 100 known extinct species with material representing the extant sister clades Strigidae (typical owls) and Tytonidae (barn owls), alongside material referred to several extinct groups of uncertain systematic affinities (i.e., Palaeoglaucidae, Necrobyinae, Selenornithinae, Sophiornithidae, Protostrigidae, and Ogygoptyngidae; reviewed by [2–4]). The relationships of these extinct groups to one another and to the crown clade members of Strigiformes has yet to be evaluated in a rigorous phylogenetic analysis. The absence of such a comprehensive systematic study of the group likely derives from the nature of the owl fossil record, with many named extinct species known only from isolated and otherwise fragmentary fossils. The oldest fossils attributed to the Strigiformes date from the late Paleocene of North America (Ogygoptynx) and Europe (Berruornis; reviewed by [3,4]). Owls evolved large body sizes early in their history, as indicated by the comparable size of specimens attributed to the presumably non-crown group owl Berruornis with that of extant Eagle Owls (Bubo spp.; [3]). The relatively high diversity of smaller sized Paleocene and Eocene fossils from North America, Europe, and Asia demonstrate that owls had diversified greatly in body-size during the Paleogene [2–4]. In contrast, pre-Quaternary strigiform fossils from Africa and the Southern Hemisphere are quite rare with a single record from the Miocene Diversity 2020, 12, 163; doi:10.3390/d12040163 www.mdpi.com/journal/diversity Diversity 2020, 12, 163 2 of 12 Diversity 2020, 12, x FOR PEER REVIEW 2 of 12 ofthe Argentina Miocene [of5] Argentina and a small [5] numberand a small of strigiform number of fossils strigiform from fossils South from Africa South [6]. Africa These [6 regionally]. These diregionallyffering fossil differing records fossil currently records obfuscate currently the reconstructionobfuscate the ofreconstruction strigiform historical of strigiform biogeography historical and thebiogeography origin of the and clade’s the orig currentin of cosmopolitanthe clade’s current geographic cosmopolitan distribution. geographic distribution. TheThe relatively relatively sparse sparse fossil fossil record record of African of African owls isowls entirely is entirely composed composed of Neogene of andNeogene Quaternary and specimensQuaternary from specimens the early from Pliocene the early of South Pliocene Africa, of South the late Africa, Pliocene the late of Morocco, Pliocene theof Morocco, early Pleistocene the early of Tanzania,Pleistocene as well of Tanzania as several, as records well fromas several Pleistocene records cave from deposits, Pleistocene and Holocene cave deposits, deposits and from Holocene islands to thedeposits east of Madagascarfrom islands [2to,7 –the14 ].east Previously, of Madagascar the oldest [2, reported7–14]. Previously, African specimen the oldest of reported Strigiformes African was a distalspecimen humerus of Strigiformes from the early was Miocene a distal (~18humerus Ma) offrom Kenya the thatearly was Miocene referred (~18 to theMa) extant of Kenya genus thatOtus was[7 ]. However,referred theto the affinities extantof genus the Kenyan Otus [7 specimen]. However, are the in need affinities of reevaluation, of the Kenyan as the specimen taxonomic are identification in need of ofreevaluation, Walker and Dykeas the [ 7taxonomic] was not supportedidentification by theof Walker description and ofDyke any strigiform[7] was not apomorphies supported by present the indescription the fossil and of theany referralstrigiform to Otus apomorphieswas subsequently present questionedin the fossil by and Pavia the et al.referral [6]. An to isolatedOtus was alar phalanxsubsequently from the questioned middle Miocene by Pavia (~13 et al. Ma) [6]. ofAn Morocco isolatedwas alar referredphalanx tofrom the the extant middle species MioceneTyto alba (~13by BrunetMa) of [ 15Morocco,16]. Naturally, was referred the potential to the extant presence species of anTyto extant alba by species Brunet in the[15,1 middle6]. Naturally, Miocene the raises potential issues relatedpresence to speciesof an extant longevity species (see in discussion the middle in [Miocene17,18]), and rais thates issues particular related taxonomic to species identification longevity (see also wasdiscussion questioned in [ by17, Pavia18]), and and that Mourer-Chauvir particular taxonomicé [19]. Therefore, identification the earliest also was reliable questioned records ofby African Pavia and owls areMourer from the-Chauviré richlyfossiliferous, [19]. Therefore, early the Pliocene earliest (~5.0 reliable Ma) depositsrecords of at African Langebaanweg, owls are South from Africathe richly [6,20 ]. Thus,fossiliferous, the discovery early of Pliocene a new strigiform (~5.0 Ma fossil) deposits from theat Langebaanweg, early Oligocene [~30South Ma] Africa Jebel Qatrani[6,20]. Thus, Formation the ofdiscovery Egypt (Figure of a 1new) extends strigiform the African fossil strigiformfrom the early fossil Oligocene record by approximately [~ 30 Ma] Jebel 25 Qatrani million years.Formation We refer of Egypt (Figure 1) extends the African strigiform fossil record by approximately 25 million years. We to this specimen (and the species it represents) herein as the ‘Jebel Owl’. refer to this specimen (and the species it represents) herein as the ‘Jebel Owl’. FigureFigure 1. 1.Maps Maps depictingdepicting thethe locationlocation of the Fayum Depression fossil fossil locality locality in in Egypt Egypt (A (A), ),Quercy Quercy fissurefissure fill fill fossil fossil owl owl localitylocality inin FranceFrance and previous African African strigiform strigiform fossil fossil localities localities (B ():B ):Quercy, Quercy, FranceFrance (1); (1); Ahl Ahl al al Oughlam Oughlam Quarry,Quarry, Casablanca,Casablanca, Morocco (2); Fayum Fayum Depression, Depression, Egypt Egypt (3); (3); Olduvai Olduvai Gorge, Tanzania (4); Mauritius and Reunion Island (5); Langebaanweg, South Africa (6). Gorge, Tanzania (4); Mauritius and Reunion Island (5); Langebaanweg, South Africa (6). Diversity 2020, 12, 163 3 of 12 2. Materials and Methods 2.1. Osteological and Geologic Conventions In the osteological description, the English equivalents of the Latin nomenclature, summarized by Baumel and Witmer [21], are used. Osteological measurements of fossil specimens and extant strigiformes follow those outlined by von den Driesch [22], and were taken using digital calipers, rounding measurements to the nearest 0.1 mm (see Supplementary Data File S1). Ages of geologic time intervals are based on the International Geologic Time Scale [23]. A complete list of extant comparative skeletal material and measurements of those specimens are included in the Supplementary Materials section. 2.2. Institutional Abbreviations CU, Campbell Museum of Natural History, Clemson University, Clemson, South Carolina, USA; DPC, Duke Lemur Center, Division of Fossil Primates, Duke University, Durham, North Carolina, USA; FMNH, Field Museum of Natural History, Chicago, Illinois, USA; MNHN, Muséum National d’Histoire Naturelle, Paris, France; NCSM, North Carolina Museum of Natural Sciences, Raleigh, North Carolina, USA; USNM, Smithsonian Institution, National Museum of Natural History, Division of Birds, Washington, District of Columbia, USA. 3. Systematic Paleontology Aves Linnaeus, 1758 [24] Strigiformes
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