A New Barn Owl (Aves: Strigidae) from the Early Miocene of Germany, with Comments on the Fossil History of the Tytoninae

Home , Deinogalerix, Gargano

J. Ornithol. 139, 247-261 (1998) © Deutsche Ornithologen-Gesellschaft/BlackwellWissenschafts-Verlag, Berlin ISSN 0021-8375

A new barn owl (Aves: Strigidae) from the early Miocene of Germany, with comments on the fossil history of the Tytoninae

JiN Ml~ovsk#

Institute of Geology and Paleontology, Charles University, Albertov 6, CZ-128 43 Praha 2, Czech Republic

Summary

A previously unknown, very large barn owl, Basityto rummeli is described from the early Miocene of Grafenmtihle 21 in Bavaria, Germany. This raises the number of tytonine genera known from the early Miocene to three. Necrobyas is known only from France, Prosybris from France and Austria, and Basityto so far only from Germany. Further taxonomic conclusions are as follows: Palaeotyto cadurcensis Mourer-Chauvirr, Palaeobyas cracrafii Mourer-Chauvirr, and Tyto edwardsi (Gaillard) are excluded from the Tytoninae. Six species are synonymized: Necrobyas rossignoli Milne-Edwards with Necrobyas harpax Milne-Ed- wards, Necrobyas edwardsi Gaillard with Necrobyas arvernensis (Milne-Edwards), Necrobyas minimus Mourer-Chauvir6 with Prosybris antiqua (Milne-Edwards), Tyto campiterrae Jfinossy with Tyto sanctialbani (Lydekker), Tyto robusta Ballmann with Tyro gigantea Ballmann, and Tyto melitensis (Lydekker) with Tyto alba (Scopoli). Necrobyas medius Mourer-Chauvir6 was transferred to the genus Prosybris. In addition, Strix ignota "Milne-Edwards" and Strix ignota "Paris" emerge as not available for nomenclatural purposes.

Key words: Aves, Strigidae, Tytoninae, Paleogene, Neogene, Quaternary

Zusammenfassung

Eine neue Schleiereule (Ayes: Strigidae) aus dem Untermioziin in Deutschland: mit Anmerkungen zur fossilen Geschichte der Tytoninae

Eine neue, sehr grof3e Schleiereule, Basityto rummeli, wird aus dem Untermioz~n yon Grafenmt~hle 21 in Bayern beschrieben. Damit ist die Anzahl der ans dem Untermioz~n bekannten Schleiereulen- g attungen auf drei gestiegen. Necrobyas ist bisher nur aus Frankreich, Prosybris aus Frankreich und Osterreich, und Basityto aus Bayern bekannt. Weitere taxonomische Ergebnisse sind wie folgt: Palaeotyto cadurcensis Mourer-Chauvir6, Palaeobyas cracrafii Mourer-Chanvir6 und Tyto edwardsi (Gaillard) wurden aus den Tytoninae entfernt. Sechs Arten wurden synonymisiert: Necrobyas ross# gnoli Milne-Edwards mit Necrobyas harpax Milne-Edwards, Necrobyas edwardsi Gaillard mit Ne- crobyas arvernensis (Milne-Edwards), Necrobyas minimus Mourer-Chauvir6 mit Prosybris antiqua (Milne-Edwards), Tyto campiterrae Jfinossy mit Tyto sanctialbani (Lydekker), Tyro robusta Ball- mann mit Tyto gigantea Ballmann und Tyto melitensis (Lydekker) mit Tyto alba (Scopoli). Necrobyas medius Mourer-Chauvir6 wurde in die Gattung Prosybris ~berffihrt. Die Namen Strix ignota ,,Milne- Edwards" und Strix ignota ,,Paris" sind ftir nomenklatorische Zwecke nicht verfiJgbar.

Introduction genera (Tyto Billberg, 1828, and Phodilus Ge- offroy-Saint-Hilaire, 1830), with 9 and 2 spe- Modem barn owls (subfamily Tytoninae of the cies (sensu Wolters 1975-1982), respectively. family Strigidae) are small to middle-sized The fossil record of the Tytoninae, which inclu- owls, which inhabit a variety ofbiotopes around des also pigmy and gigantic forms, is quite rich the world (Burton 1973, Eck & Busse 1973, and goes back to the late Eocene (Mourer-Chau- Schneider 1977). The subfamily (treated as fa- vir6 1987). mily by some authors) consists now only of two In the present paper I will describe a new barn

U.S. Copyright Clearance Center Code Statement: 0021-8375/98/13903-0247 $ 11.00/0 248 Journal ftir Ornithologie 139, 1998 owl species from the early Miocene of Bavaria, Germany, and comment on the fossil history of the subfamily Tytoninae. The stratigraphy follows Schmidt-Kittler (1987) for the Paleogene (Mammal Paleogene zones, MP), Mein (1990) for the Neogene (Mam- mal Neogene zones, MN), and Horfi6ek & Lo~ek (1988) for the Quaternary (Quaternary biozones, Q). Museum acronyms are as follows: MNHN = Mus6um National d'Histoire Naturelle, Paris, France; NHMW = Namrhistorisches Museum, Wien, Austria; and USNM = United States Na- tional Museum, Washington, D.C., USA.

Systematic Paleontology

Order Strigiformes Wagler, 1830 Family Strigidae Vigors, 1825 Subfamily Tytoninae Ridgway, 1914

Basityto, n. g. Type: Basityto rummeli, n. sp. Included species: Type species only. Diagnosis: Very large barn owl. Humerus with high ectepicondyle, distal margin between exter- b nal and internal condyle slightly deepened, ligamental furrow broad, deep, almost perpendicu- Fig. 1. Holotype humerus of Basityto rummeli from lar to the axis of the bone, and distally opened. the early Miocene of Bavaria (a), and humerus of a modern Barn Owl Tyro alba (b) in anconal view Comparison: Holotypical humerus of Basi- (55% of natural size) tyto rummeli agrees with the same element of Abb. 1. Der holotypische Humerus yon Basityto the Tytoninae, and differs from that of other rummeli aus dem UntermiozS.n Bayerns (a) und Hu- owls, in having a large and long ligamental merus einer rezenten Schleiereule Tyro alba (b) in furrow. It differs from the humeri of Necrobyas Medialansicht (55% der natiMichen GrSge) in having a higher ectepicondyle, and a slightly deepened distal margin between internal and owl, which virtually made it the feathered king external condylus; from the humeri of Noctur- of Central European early Miocene. navis in having a much broader ligamental fur- Basityto rummeli, n. sp. (Fig. la, 2) row perpendicular to the axis of the bone; from Tyto and Phodilus in having a higher ectepicon- Holotype: Almost complete left humerus in two dyle, and a much broader ligamental furrow parts; coll. Rummel, uncatalogued. perpendicular to the axis of the bone. No direct Material: Holotype only. comparison was possible with the humeri of Age and locality: Early Miocene, MN 2-3, of Prosybris and Selenornis, because this element Grafenmt~hle 21, Bavaria, Germany (M. Rum- is not known for these two genera. Neverthe- mel, in litt.). The locality is one of the many less, Prosybris was a genus of pigmy owls, and fossiliferous fissures in southern Germany (see Selenornis was also much smaller than Basi- Ml~kovsk~ 1992, Ml~ovsk~) & Hesse 1996). tyro. Etymology: Abbreviated from Greek Diagnosis: As for the genus. [3c~o-t)~Evg, king, and Tyto, modem genus of Measurements: Greatest length (estimated) = barn owls. Alludes to the majestic size of the 175 ram, greatest width of distal end = ca. J. Mlikovsk)~.A new barn owl from the early Miocene 249

Comments on some other fossil barn owls

Brodkorb (1971) listed 11 species of barn owl, to which Mourer-Chauvir6 (1987) added an- other ten. Below I present conunents on some of these taxa.

Palaeotyto cadurcensis PaIaeotyto cadurcensis was described by Mourer-Chauvir6 (1987) in the monotypic genus Palaeotyto on the basis of a coracoid from an unknown locality within the Phos- phorites du Quercy. Age of the species is unknown, and can lie anywhere between the middle Eocene and late Oligocene (R6my et al. 1987, Mourer-Chauvir6 1995, 1996). The coracoid of Palaeotyto differs from the same element of proper Tytoninae in the configuration of its head, and in the remarkable size of the coracoidal foramen. For further details of how the coracoid of Palaeotyto differs from the same element of Necrobyas, Tyto and Phodilus, the three most important barn owl genera, see Mourer-Chauvir6 (1987). Consequently, Pa- laeotyto should be removed from the Tytoni- nae. Fig. 2. Holotype humerus of Basityto rummeli from the early Miocene of Bavaria in plantar view (55% Palaeobyas cracrafli of natural size) Palaeobyas cracrafii, the only species included Abb. 2. Der holotypische Humerus von Basityto in was described by Mourer-Chau- rummeli aus dem Untermioz~inBayerns in Lateralan- Palaeobyas, sicht (55% tier nattirlichen Gr6f3e) vir6 (1987) on the basis of a single tarsometatarsus from an unknown locality within the 26.5 ram, width x depth of shaft in the center = Phosphorites du Quercy, which range in age 13.1 x 11.2 ram. from the middle Eocene to the late Oligocene (R6my et al. 1987, Mourer-Chauvir6 1995, Etymology: After Dr. Michael Rummel 1996). The tarsometatarsus is very stout, troch- (Weissenburg), who collected the fossil, in rec- leae are open in distal view, and external hypo- ognition of his paleontological work on verte- tarsal ridge is blunt. In these features Palae- brate remains from the Tertiary fissure deposits obyas differs from the Tytoninae, and agrees of Bavaria. with Sophiornis and Berruornis, which are Remarks: Basityto rummeli is the largest con- placed in the family Sophiomithidae (Mourer- tinental barn owl known, being comparable in Chauvir6 1987, 1994). Hence, Palaeobyas size to the gigantic island forms Tyto riveroi should be removed from the Tytoninae, and A~edondo, 1972b from the Quaternary of placed in the Sophiornithidae. Cuba, and Tyto gigantea Ballmann, 1973 from the Pliocene of Gargano. It probably preyed Necrobyas spp. upon small ungulates, which are abundant in Mourer-Chauvir6 (1987) distinguished six south German fissure deposits, formed during species in the genus Necrobyas: N. harpax the time when Basityto rummeli inhabited the Milne-Edwards, 1892, N. rossignoli Milne-Ed- region (see Heissig 1978). wards, 1892, N. edwardsi Gaillard, 1939, 250 Journal ftir Ornithologie 139, 1998

N. medius Mourer-Chauvir6, 1987, N. minimus should be deemed to be the holotype of Necro- Mourer-Chauvir6, 1987, and N. arvernensis byas harpax. (Milne-Edwards, 1863). Of these, Neerobyas Necrobyas rossignoli was based by Milne- medius and Necrobyas minimus differ distinctly Edwards (1892: 63) on a single tarsometatarsus from Neerobyas in the morphology of their whose exact age he neglected to mention, from tarsometatarsi, and belong in the genus Prosy- the deposits of the Phosphorites du Quercy, bris (see below). which range in age between the middle Eocene Before addressing the taxonomic status of the and late Oligocene (Mourer-Chauvir4 1995, remaining fo.ur species, three technical com- 1996, R4my et al. 1987). Mourer-Chauvir6 ments are necessary: Necrobyas harpax was (1987) decided that the species was late Eocene based by Milne-Edwards (1892) on a tarsome- in age. This observation was based on a single, tatarsus, to which an ulna and a humerus were very fragmentary coracoid, of which only the assigned ("Je rapporte ~..."). Hence, the tar- shaft was measurable, which came from the late sometatarsus is the holotype, while ulna and Eocene (MP 17) of Perri~re in Quercy. Metrical humerus are paratypes of the species. Inexplic- comparison (Fig. 3) shows that this specimen ably, Mourer-Chauvir6 (1987: 97) selected left differs from Oligocene coracoids of Necrobyas tarsometatarsus (MNHN QU 15695) as a lecto- spp. in the shape of its shaft. Hence, I doubt that type (sic!) of Necrobyas harpax. This action the specimen was correctly assigned to Necro- has no bearing on the nomenclatural or taxo- byas, let alone Necrobyas rossignoli. In conse- nomic Status of the species. Moreover, meas- quence, there is no indication that Necrobyas urements of this "lectotype" differ markedly rossignoli is a late Eocene species. Similarly, from the measurements of the holotype given there is no indication that the genus Necrobyas by Mitne-Edwargs (1892: 62), which are as existed before the Oligocene. follows: maximum length = 37 ram, proximal Necrobyas arvernensis was described by width = 8.5 mm, width of the shaft = 5 mm, and Milne-Edwards (1863) on the basis of a tar- distal width= l0 ram. Among Necrobyas tarsometatarsus and a tibiotarsus from the early sometatarsi from early collections, which are Miocene (MN 2) of Saint-G6rand-le-Puy in deposited in MNHN, and could thus be at the France. Most fossil owls were based on tar- disposal~ofMilne-Edwards, only the tarsometa- sometatarsi, while tibiotarsi are much less tarsus QU 15742 closely resembles Milne-Ed- abundant. Hence, I select here the tarsometa- wards" 'specimen in measurements (see tarsus MNHN Av. 2834b as the lectotype of Mourer-Chauvir6 1987: 102). This specimen Necrobyas arvernensis. Herewith, the tibio-

2.6

2.5

2.4 • Q

E 2.3 Fig. 3. Dimensions of Necroby- as coracoids, showing width 2.2 and depth of shaft. Data are from ,.,I--- 0_ Mourer-Chauvir6 (1987). Circle 2.1 marks the coracoid from Petrie- £3 re. See text for explanation 2 Abb. 3. Breite und Tiefe des Coracoidk6rpers der Gatmng 1.9 Necrobyas. Die Angaben ent- starnmen Mourer-Chauvir6 1.8 i i f f i i (1987). Der Kreis bezeichnet 2.6 2.8 3.0 3.2 3.4 3.6 3.8 4,0 4,2 4.4 4.6 das Coracoid aus Perri~re. Sie- Width [mm] he Text ftir Erkl~-ung J. Ml~ovsks~ • A new barn owl from the early Miocene 251

Fig. 4, Dimensions of Necrobyas 10.5 tarsometatarsi, showing their maximum length and proximal width. Data are from Mourer- 10.o Chauvir6 (1987). 1 - holotype of -~ g Necrobyas rossignoli, 2 - holoty- E 9.5 pe of Necrobyas harpax, 3 - holotype of Necrobyas edwardsi, 4 ~3 - lectotype of Necrobyas arver- "~ 9.0 • 02 nensis -d 01 Abb. 4. Maximale L~nge and .~ 8.5 proximale Breite der Tarsometa- o tarsi von Necrobyas. Die Anga- ca_ ben entstammen Mourer-Chauvi- 8.o r6 (1987). 1 - Holotyp von Ne- crobyas rossignoli, 2 - Holotyp yon Necrobyas harpax, 3 - Holo- 7.5 typ von Necrobyas edwardsi, 4 - 30 32 34 36 38 40 42 44 46 Lektotyp von Necrobyas arver- Maximum length [mm] nensis Fig. 5. Dimensions of Necroby- 12.0. as tarsometatarsi, showing their 4 o maximum length and distal 1T.5. width. Data are from Mourer- 3 • o Chauvir6 (1987). 1 - holotype 11.o ofNecrobyas rossignoli, 2 - holotype of Necrobyas harpax, 3 - E holotype of Necrobyas edward- x: 10.5 • 2 si, 4 - lectotype of Necrobyas o arvernensis. "'~ 10.o ABB. 5, Maximale L~nge and if_ distale Breite der Tarsometatar- 2"C~ si von Necrobyas. Die Angaben 9.5. entstammen Mourer-Chauvir6 1 (1987). 1 - Holotyp von Necro- 9.0. go byas rossignoli, 2 - Holotyp von Necrobyas harpax, 3 - Ho- lotyp yon Necrobyas edwardsi, 8.5 4 - Lektotyp von Necrobyas ar- 30 32 34 36 38 40 42 44 46 vernensis. Maximum length [mm]

tarsus MNHN Av. 2834a becomes the paralec- crobyas harpax and Necrobyas arvernensis, re- totype of the species. spectively. Accordingly, I synonymize here Ne- All Necrobyas species were based on tar- crobyas rossignoli Milne-Edwards, 1892 with sometatarsi, and this element is by far the best Necrobyas harpax Milne-Edwards, 1892, and represented in the collections. Metrical com- Necrobyas edwardsi Gaillard, 1939 with Ne- parisons (Fig. 4 and 5) show that these tarsome- crobyas arvernensis Milne-Edwards, 1863. Ne- tatarsi fall into two distinct groups which differ crobyas harpax was recorded only from the in size, but not in shape. The group of smaller early Oligocene (MP 21-23), while Necrobyas tarsometatarsi includes holotypes of Necrobyas arvernensis from the late Oligocene to the early harpax and Necrobyas rossignoli, while the Miocene (MP 28 - MN 2) of France. Sub- group of larger tarsometatarsi includes the ho- sequent research may show that Necrobyas har- lotype ofNecrobyas edwardsi and the lectotype pax and Necrobyas arvernensis are chronos- of Necrobyas arvernensis. These two groups pecies from a single phylogenetic lineage, ff so, can be interpreted as paleospecies (Mlikovsk~ all these forms should be treated as a single et al. 1985), which should bear the names Ne- species (cf. Haffer 1995). 252 Journal ftir Ornithologie 139, 1998

Prosybris spp. from Oligocene deposits of Fonbonne 1 in Quercy, for which no closer datation is avail- The genus Prosybris was created by Brodkorb able. It agrees in size and shape with the same (1970) for Strix [= Tyto] antiqua, described by element of Prosybris antiqua from the early Milne-Edwards (1869: 498) on the basis of a Miocene (MN 2) of France. Hence, I sy- tarsometatarsus from the early Miocene (MN 2) nonymize here Necrobyas minimus Mourer- of Saint-Grrand-le-Puy in France. The absence Chauvir6, 1987 with Strix [= Prosybris] anti- of an ossified supratendineal bridge clearly in- qua Miine-Edwards, 1869. dicates that Prosybris was a barn owl which Prosybris medius was larger than Prosybris differed from Tyto in having shorter and more antiqua, and represents a second species of the robust tarsometatarsi, and from Necrobyas in genus. Unfortunately, its age is unknown, being having slender and less robust tarsometatarsi. anywhere between the middle Eocene and late These three genera differ also in the proportions Oligocene (cf. Rrmy et al. 1987, Mourer-Chau- of their hind limbs, the relative lengths of vir6 1995, 1996). Mourer-Chauvir6 (1987) as- femur, tihiotarsus and tarsometatarsus being signed a distal end of a tibiotarsus from the approximately 1:1.7:1.2 in Tyto, 1:1.7:1.0 in early Oligocene (MP 23) of Itardies in Quercy Prosybris, and 1:1.3:0.7 in Necrobyas. to her Necrobyas medius. The specimen is se- A flattened partial skeleton of a small bird verely damaged and not suitable for exact ident- from the early Miocene (MN 3-4) of Limberg ification, although there is evidence that it has in Lower Austria was described by Bachmayer originated from a small owl (see Mourer-Chau- (1980) as that of a falcon, but my reexamination vir6 1987, pl. 2, fig. 21-22). There is thus no of the specimen (NHMW 1977/1913) showed proof that Prosybris already existed in the early that it is a typical barn owl. Its tarsometatarsus Oligocene (contra Mourer-Chauvir6 1987). agrees in size and shape with the same element Prosybris antiqua and Prosybris medius of Prosybris antiqua, so that I assign here the were pigmy barn owls, which have so far been specimen to this species. The measurements of recorded from the Oligocene (MP?) to the early the Limberg specimen are as follows: skull: Miocene (MN 3-4) of France and Austria. length = 42 mm, width (flattened) = 33 rnm; femur: greatest length = 33 ram, proximal Tyto sanctialbani width = 6.5 ram, distal width = 8.5 ram; tibiotarsus: greatest length = 55 man, proximal width The modem genus Tyto appeared in Europe in = 5.5 ram, distal width = 5 mm, width of shaft the middle Miocene (MN 7). Miocene and = 3 ram; tarsometatarsus: greatest length = Pliocene bones of Tyto, which were found in 34 ram, proximal depth = 6.5 ram. Exact meas- European deposits, fall into three groups, which urements could not be obtained, the presented can be identified as species (M1Novsk2~ et al. values are thus rounded to 1 mm (lengths) and 1985). They should bear the following names: 0.5 mm (widths and depth), respectively. Note Tyto sanctialbani (Lydekker), Tyto balearica that these values differ more or less from those Mourer-Chauvir6 et al., and Tyto gigantea Ball- given by Bachmayer (1980), because he, appar- marln. ently misled by the supposed similarity of the Tyto sanctialbani was described by Lydekker fossil with falcons, misinterpreted the shape (1893) on the basis ~f nine bones from the and size of its bone ends. middle Miocene (MN 7-8) of La Grive-Saint- Necrobyas minimus and Necrobyas medius Alban in the Department of Is~re, France. Fur- were described by Mourer-Chauvir6 (1987) on ther material of this species from the same com- the basis of distal parts of two tarsometatarsi plex locality was described by Ballmann from the Phosphorites du Quercy. These two (1969a). In the U.S. National Museum I identi- tarsometatarsi differ from the same element of fied additional material of this species from its Necrobyas and agree with that of Prosybris in type locality. The measurements are as follows: having its shaft more slender, and its external proximal width of left carpometacarpus trochlea more flaring and more distant from the (USNM 205291) = 10.0 ram, distal width of left distal end of the medial trochlea. The holotype tibiotarsus (USNM 187665) = ca. 8.8 mm, tarsometatarsus of Necrobyas minimus came proximal width of left tarsometatarsus (USNM J. M1Novsk~ • A new barn owl from the early Miocene 253

2170) = ca. 9 ram, distal width of left tarsome- bones fall in the same size class. In the absence tatarsi (USNM 187666 and 187667) = 10.8 rnm of morphological differences I synonymize and 10.9 ram. here Tyto campiterrae J~nossy, 1991 with Strix I found further bones, which are clearly refer- [= Tyto] sanctialbani Lydekker, 1893. able to Tyto sanctialbani, in the material from Ballmann (1973, 1976) reported on the rec- the late Miocene (MN 10) of Kohfidisch in ord of Tyto sanctialbani from the Pliocene is- Burgenland, Austria (see Mh'kovsk~ 1996a for land (now peninsula) of Gargano in Italy (see details on the locality). The material (deposited Delle Cave 1996 for the age of the locality). The in NHMW) includes: fragments of left and right bones (partial coracoid, ulna, tibiotarsus, tar- coracoids (not measurable), the proximal end of sometatarsus, and phalanges digitomm pedis) a left tibiotarsus (width = 10.1 ram), and the are larger and more robust than the same ele- proximal end of a left tarsometatarsus (width = ments of Tyto sanctialbani (see measurements 10.1 ram). in Ballmann 1973, 1976), in which feature they Tyto campiterrae JS.nossy, 1991 was based agree with the same elements of Tyto balearica. There is no doubt that they should be referred on a left tarsometatarsus from the late Miocene to the latter species. (MN 13) of PolgS.rdi 5 in Hungary. The only difference between Tyto campiterrae and Tyto The evidence presented above extends both sanctialbani mentioned by J4nossy (1991) was the temporal and geographic distribution of the smaller size of the former species. Unfortu- Tyto sanctialbani. The species is now known nately, he used for comparison measurements from the middle Miocene to the late Miocene of of a specimen from San Giovannino in Italy La Grive-Saint-Alban in France (MN 7-8), Kohfidisch in Austria (MN 10), and Polg4rdi in (following Ballmann 1973), which actually be- Hungary (MN 13). Tyto sanctialbani was a barn longs to Tyto balearica (see below). Metrical owl of the size of the modem Tyto alba, and comparisons of the holotype tarsometatarsus of with similarly slender bones. Tyro campiterrae with the same element of Tyto sanctialbani are as follows: proximal width = Tyro balearica 10.2-11.0 mm (n = 8) vs. ca. 9-10.1 mm (n = 2), distal width = 11.2-11.7 mm (n = 5) vs. Tyto balearica was described from the latest 10.8-10.9 rnm (n = 2) in Tyro campiterrae and Pliocene (MN 17) of Cova de Canet on the Tyto sanctialbani, respectively. Hence, all these island of Mallorca (Mourer-Chauvir6 et al.

g' !,. //

Fig. 6. Distribution of Tyro sanctialbani C~) and Tyro balearica (I) from the middle Miocene to the middle Pleistocene Abb. 6. Verbreitung yon Tyto balearica ~) und Tyto balearica (B) im Zeit- raum von Mittelmioz~n bis Mittelpleistoz~n 254 Journal ftir Omithologie 139, 1998

1980). Subsequently it was found also in sev- learica. This error is understandable, because eral late Pliocene to early Pleistocene localities Ballmann (1973, 1976) befieved that the island on the nearby mainland in Spain and France was inhabited by slender-boned Tyro sanctial- (Mourer-Chauvir6 & Sanchez Marco 1988), in bani, whereas it was inhabited by a more robust the late Miocene of Aljezar B in Spain (Chene- Tyto balearica, the existence of which was not val & Adrover 1995), and in the middle Pleis- yet recognized in the mid 1970s. tocene of Castiglione 3 in Corsica (C. Mourer- The Pliocene history of barn owls on Gar- Chauvir6 in Salotti et al. 1997). In addition, gano can be summarized as follows: originally, bones from the Pliocene of Gargano, assigned the island was inhabited by Tyto balearica, by Ballmann (1973, 1976) to Tyto sanctiaIbani, which was widespread in the western Mediter- belong here (see above). ranean in that time (Fig. 6). Rodents and insec- Tyto balearica was slightly larger than both tivores, which presumably was the main food of Tyto sanctialbani and Tyro alba, and its bones Tyro balearica on Gargano, evolved toward were markedly more robust. Its known distribu- larger body size (see Freudenthal 1971, 1972, tion ranges from the late Miocene (MN 12) to 1976), which forced their predator to increase the middle Pleistocene (Q 3) of Spain, France its body size as well. At that time, the island was and Italy (Fig. 6). inhabited by but a single barn owl species, to which Ballmann (1973, 1976) applied the name Tyro gigantea Tyto robusta. Continuing increase in body size allowed subsequently Tyto balearica to re- Ballmann (1973) described from the Neogene settle on the island. Interspecific competition island (now peninsula) of Gargano two new between Tyro balearica and Tyto robusta species of large barn owls: Tyto robusta, and forced the latter species to further increase its Tyto gigantea. The "locality" is a complex of body size. The latter form, known only from fissure deposits which differ in age, but all were youngest deposits was named by Ballmann believed to belong in the late Miocene, when (1973, 1976) Tyto gigantea. There is no evi- Ballmann (1973, 1976) described the bones dence for the contemporaneous existence of (Freudenthal 1971, 1976). However, sub- robusta and gigantea (see above; contra Ball- sequent research made it much more probable mann 1973, 1976). Taking into account that that the Gargano fauna is early Pliocene in age Gargano was a small island when these barn (see Delle Cave 1996). owls lived there, robusta, and gigantea can be Ballmann (1973, 1976) showed that the is- interpreted as temporary representatives of a land was originally inhabited by a single barn single lineage of barn owls, evolving toward owl species, erroneously identified by himself larger size. Such an evolution has been well as Tyto sanctilabani (= balearica; see above). documented on many Quarternary islands (see In younger deposits, the size of the barn owls below). I agree with Haffer (1995) that the artifi- from Gargano increased (Ballmann's robusta), cial delimitation of temporal portions of a lineage but small (balearica) and very large (gigantea) as chronospecies is meaningless. Hence, I sy- forms were absent. In youngest deposits, the nonymize here Tyto robusta Ballmarm, 1973 with size of the barn owls from Gargano ranged from Tyto gigantea Ballmann, 1973. small to very large. Ballmann (1973, 1976) interpreted the latter observation as evidence of Tyro melitensis the contemporaneous presence of three barn owl species on Gargano during that time. Tyto melitensis was described by Lydekker Nevertheless, examination of fig. 8 in Ball- (1891) on the basis of a femur from the Quater- mann (1976: 23) clearly shows that the size of nary (middle Pleistocene) deposits of Malta in barn owl bones from youngest deposits (par- the belief that it is "slightly longer and more ticularly from San Giovannino) fall into two slender" than the same element of modern Tyto clusters. Bones from youngest deposits, identi- alba, which it is not, as was already observed fied by Ballmann (1973, 1976) as robusta, be- by Mourer-Chauvir6 et al. (1980). According to longed in fact to large individuals of Tyto ba- Lydekker (1891) length of the holotype femur J. Mlikovsk~ • A new barn owl from the early Miocene 255 of Tyto melitensis is 54 ram, while it is 48.0- brecht (1921: 97) inexplicably stated that Strix 54.3 mm in European (n = 28, Cheneval & ignota Milne-Edwards is based on the distal Adrover 1995, Mlfkovsk3~ unpub, data), 52.0- portion of a tarsometatarsus from the middle 55.1 mm in African (n -- 2), 51.4-56.1 mm in Miocene of Sansan in France, figured in Milne- Asian (n = 2), and 58.5-63.4 mm in North Edwards (1869, pl. 192, fig. 1-2). This spe- American Tyto alba (n = 11). Width of the shaft cimen was referred by Milne-Edwards (1869: is 4 mm in Tyto melitensis (Lykker 1891), while 499) to "Strix sp.", and the name Strix ignota the corresponding dimension in the European does not appear even in the legend to Milne-Ed- Tyto alba is 3.9-4.3 mm (n = 28, Cheneval & wards' plate No. 192. Later, Lambrecht (1933: Adrover 1995, Mlikovsk2~ unpub, data). There 613) attributed the name to Paris himself, stat- is thus no reason to separate Maltese barn owls ing that Paris applied this name to Strix sp. of at the species level, and I synonymize here Strix Milne-Edwards ("1871": 499). Nevertheless, [= Tyrol melitensis Lydekker, 1891 with the Lambrecht's (1933) approach cannot be ac- modem Tyto alba (Scopoli, 1769). cepted. Milne-Edwards (1869) devoted the upper part of his p. 499 to Strix antiqua (this Strix edwardsi chapter started on p. 498), and its lower part to Strix sp. Both of these taxa are correctly listed Strix [= Tyto] edwardsi was described by En- by Paris (1912: 287), which rules out the possi- nouchi (1930) from the middle Miocene (MN bility that he applied Strix ignota to Milne-Ed- 7-8) of La Grive-Saint-Alban in France on the wards' Strix sp. (as believed by Larnbrecht basis of distal part of a tibiotarsus, which was 1933, and Brodkorb 1971). Moreover, Paris said to be smaller than the same element of Strix (1912) labeled Strix ignota with Milne-Ed- [= Tyto] sanctialbani, described by Lydekker wards' name, which indicates that he did not (1893) from the same locality, the distal width intend to name the species. Brodkorb (1971: of tibiotarsus being 10.6 mm in Tyto sanctial- 230) inexplicably stated that Strix ignota was bani, and 7.2 mm in the holotype of Tyto ed- created by Paris (1912) as a new name (!) for wardsi, respectively (Ennouchi 1930). As Strix sp. Milne-Edwards, 1871 [= 1869]. The judged from the figures in Ennouchi (1930, pl. latter does not include any species-group name, V., figs. 9-12), the holotype of Strix edwardsi however, so that Brodkorb's interpretation is differs from the same element of the Tytoninae, meaningless. Sunmaarizing this evidence, it and agrees with that of the Striginae, in having: seems probable, that Paris (1912) simply listed (1) transition between posterior margin of con- Strix ignota in error. All in all, Strix ignota dyli and the shaft smooth, (2) internal condylus Milne-Edwards never did exist, and the name is flattened, (3) anterior intercondylar fossa deep, not available for nomenclatural purposes with and (4) posterior intercondylar fossa deep. either Milne-Edwards' or Paris' name. Hence, the species must be removed from the Tytoninae in the Striginae. Strigine owls were Varia abundant and diverse in the Miocene of Europe, but are in need of revision (Mlfkovsk3) 1996b). Genus Palaeoglaux with a single species Pa- Hence, I will not attempt here to fix the taxo- laeoglaux perrierensis from the late Eocene of nomic position of Strix edwardsi within the Perri~re in France, which was described by Striginae. Mourer-Chanvir6 (1987) as a barn owl, was removed from the family by Peters (1992). Strix ignotus Ml~ovsk2~ (1996b: 809) erroneously listed a Paris (1912: 287) and Lambrecht (1921: 97) barn owl record from the Oligocene of Libya. listed among the fossil barn owls of France also The record (a fragmentary humerus from Jebel Strix [= Tyto] ignotus Milne-Edwards, 1871 Zelten) refers to a stork (see Mlikovsk~, in [= 1869], referring to p. 499 of Milne-Edwards' press). The alleged barn owl record from Libya book. However, no such name appears on that thus should be deleted from literature. page, nor elsewhere in Milne-Edwards' Alleged barn owls Strix [= Tyto] sauzieri treatise, which caused much confusion. Lain- Newton & Gadow, 1893 and Strix [= Tyto] 256 Journal ftir Omithologie 139, 1998

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~. ~ = o !I ~i ~ ~u 258 Journal ftir Omithologie 139, 1998 newtoni Rothschild, 1907 from the subfossil barn owls were diverse in Europe (Tab. 3). The deposits of Mauritius in the Indian Ocean were taxa included Necrobyas harpax (early Off- shown to belong in a single species, Masca- gocene of France), Necrobyas arvernensis (late renotus sauzieri, closely related to Otus Oligocene to early Miocene of France), Prosy- (Mourer-Chanvir6 et al. 1994). bris antiqua (Oligocene to early Miocene of In describing the barn owl Lechuza stirtoni, France and Austria), and Basityto rummeli Miller (1956) believed that its holotypical co- (early Miocene of Germany). In addition, Sele- racoid came from the middle Pliocene of Cali- nornis henrici and Prosybris media, which fornia. Chandler (1982) showed that the bone is were described from undated (middle Eocene to not fossil, and that it is inseparable from the late Oligocene) deposits of the Phosphorites du modem Tyto alba. In consequence, he sy- Quercy in France (Mourer-Chauvir6 1987), nonymized Lechuza stirtoni Miller with Tyro could belong to this time period. alba (Scopoli). Barn owls disappeared from Europe by the end of the early Miocene (none were recorded Fossil history of the Tytoninae in MN-zones 5-6, and even the record from MN 4 is uncertain), to return in MN 7 with the Taking into account all the taxonomic changes modem genus Tyro. It is notable that this extinc- mentioned above, it is possible to summarize tion of barn owls coincides in Europe with the here the available evidence of the fossil history appearance of strigine owls. Their first record of the Tytoninae as follows. comes from the early Miocene (MN 2) of Saint- Although the record of owls goes back to the Grrand-le-Puy in France (Milne-Edwards Paleocene (Rich & Bohaska 1976, 1981, 1863, 1869). Soon, these owls became more Mourer-Chauvir6 1994), the oldest owl prob- diversified, as evidenced by records from MN ably referable to the Tytoninae (Tytonidae 3 localities Wintershof (West) in Germany (Ballmann 1969a), and Merkur in the Czech auct.) is Nocturnavis incerta from the late Eocene (MP 19) of Escamps in France Republic (M1Novsk2~, unpub, data). Competi- tion with the advanced Striginae could thus be (Mourer-Chauvir6 1987). This is the only rec- the reason why tytonine owls received such a ord of the Tytoninae sensu stricto before the set-back towards the end of the early Miocene "grande coupure" at the Eocene/Oligocene (in Europe at least). boundary, which markedly separates both avian and non-avian faunas in Europe (Pomerol & From the middle Miocene onwards, only Premoli-Silva 1986, Mh'kovsk~ 1996b). barn owls of the modem genus Tyto are known During the Oligocene and early Miocene, from Europe. The species recognized include

Table 3. Stratigraphical distribution of the Tytoninae, MP and MN zones were lumped into standard mammal ages (Schmidt-Kittler 1987, Mein 1990) Tab. 3. Stratigraphische Verbreitung der Tytoninae. MP- und MN-Zonen wurden in standardisierte S~iugetie- repochen vereinigt (Schmidt-Kittler 1987, Mein 1990) MP MN Q

14-16 17-20 21-24 25-30 1-2 3-5 6-8 9 10 11-13 14-15 16 17 1-2 3-4 Selenornis ? ? ? ? Nocturnavis Necrobyas Prosybris Basityto Tyro Genus indet. I Phodilus l t" J. Mh'kovskS,. A new barn owl from the early Miocene 259

Tyto sanctialbani (MN 7-13 of France and Ger- References many), Tyto balearica (MN 12-Q3 of Spain, France and Italy), and Tyro gigantea (early? Arredondo, O. (1972a): Nueva especie de ave f6sil Pliocene of Gargano island in the Adriatic Sea). (Strigiformes: Tytonidae) del Pleistoceno superior In addition, an undescribed barn owl was found de Cuba. Bol. Soc. venezol. Cienc. nat. 29: 415- 431. in the deposits of Love Bone Bed in Horida Arredondo, O. (1972b): Especie nueva de lechuza (Becker 1987), which correspond in age to MN (Strigiformes: Tytonidae) del Pleistoceno cubano. 10. This is the oldest (and the only Tertiary) Bol. Soc. venezol. Cienc. nat. 30: 129-140. record of a tytonine owl outside of Europe. Arredondo, O. (1976): The great predatory birds of In Quaternary continental deposits, only the Pleistocene of Cuba. In: Olson, S.L. (ed.): modern species of the genus Tyto have been Collected papers in avian paleontology honoring the 90th birthday of Alexander Wetmore. Smith- found (Lambrecht 1933, Brodkorb 1971, and son. Contrib. Paleobiol. 27: 169-187. more recent studies), starting with Tyto alba Bachmayer, F. (1980): Ein fossiler Vogelrest aus den from the earliest Pleistocene (Q 1) of Oldovai Diatomeen-Schiefern (Mioz~, Ottnangien) yon Gorge in Tanzania (Brodkorb & Mourer-Chau- Limberg, Niederrsterreich. Ann. naturhist. Mus. vir6 1984). On the other hand, various endemic Wien 83: 25-28. Tyto species, sometimes rather large, have been Ballmann, P. (1969a): Die VOgel ans der altburdiga- described from several oceanic and epicon- len Spaltenfiillung von Wintershof (West) bei Eichstiitt in Bayern. Zitteliana 1: 5-61. tinental islands, including New Caledonia (Tyto Ballmann, P. (1969b): Les oiseaux mioc~nes de La letocarti Balouet and Olson, 1989), Puerto Rico Grive-Saint-Alban (Is~re). Gdobios 2: 157-204. (Tyro cavatica Wetmore, 1920), Haiti (Tyto os- Ballmann, P. (1973): Fossile VOgel aus dem Neogen tologa Wetmore, 1922; see also Olson & Hil- der Halbinsel Gargano (Italien). Scripta geol. 17: gartner 1982), Bahamas (Tytopollens Wet- 1-75. more, 1937; see also Olson & Hilgartner 1982), Ballmann, P. (1976): Fossile V6gel ans dem Neogen der Halbinsel Gargano (ltalien), zweiter Teil. and Cuba (Tyro noeli Arredondo, 1972a, and Scripta geol. 38: 1-59. Tyro riveroi Arredondo, 1972b; see also Arre- Balouet, J.C. & OIson, S.L. (1989): Fossil birds from dondo 1976). late Quaternary deposits in New Caledonia. Smith- Today, only two genera of barn owls survive. son. Contrib. Zool. 469: 1-38. Tyto is ubiquitous, while Phodilus is limited to B ecker, J.J. (1987): Neogene avian localities of North America. Washington, D.C. the forests of central Africa and south-east Asia. Brodkorb, P. (1970): Two fossil owls from the Aqni- Morphologically, Phodilus bears many features tanian of France. Quart. J. Florida Acad. Sci. 32: of the Paleogene barn owls (cf. Mourer-Chau- 159-160. vir6 1987) and it can be hypothesized that it is Brodkorb, P. (1971): Catalogue of fossil birds: Part4 phylogenetically nearer to these owls than to (Columbiformes through Piciformes). Bull. Flori- Tyto. Its antiquity is sflpported also by its dis- da State Mus. (Biol. Sci.) 15: 163-266. junctive neoendemic distribution. Brodk0rb, P. & Mourer-Chauvirr, C. (1984): Fossil owls from early Man sites of Olduvai Gorge, Tan- zania. Ostrich 54: 17-27. Burton, J.A. (ed., 1973): Owls of the world. New York. Acknowledgements Chandler, R.M. (1982): A reevaluation of the Plioce- The barn owl humerus from Grafenm/fialewas placed at my ne owI Lechusa st#toni Miller. Auk 99: 580-581. disposal by its collector Michael Rummel (Weissenburg). In Cheneval, J. & Adrover, R. ("1993" = 1995): L'avi- addition, I was allowed to~tudy bones of fossil barn owls in faune du Miocene suprrieur d'Aljezar B (Los A1- the Namrhistorisches Museum in Wien, Austria (Heinz Koll- jezares, Province de Teruel, Espagne). Systrmati- mann, Ortwin Schultz & Gudrun Daxner-Hrck), and in the que et palrodcologie. Paleontologia i Evoluci6 26- United States National Museum in Washio4gt~n,D.C. (Storrs 27: 133-144. L. Olson). The drawings were prepared by Jan Hogek (Tmo- vfi). Comments of two anonymous referees helped to clarify Delle Cave, L. (1996): Tertiary avian localities of the text. My thanks are due to all-of-these.people.My studies Italy. In: Ml~ovsk~, J. (ed.): Tertiary avian locali- in the United States National Museum in Washington, D.C. ties of Europe. Acta Univ. Carol. (Geologica) 39: were conducted when I was a short-term fellow of the 665-681. Smithsonian Institution in January/February 1997. My trips Eck, S. & Busse, H. (1973): Eulen (Aves, Strigidae). to Vienna were supported bythe Namrhistorisches Museum Wittenberg Lutherstadt. Wien. Ennouchi, E. (1930): Contribution 5 l'~tude de la 260 Journal for Omithologie 139, 1998

faune du Tortonien de la Grive-Saint-Alban (Is~- M1Novsk3), J. (1996a): Tertiary avian localities of re). Paris. Austria. In: Ml~ovsky, J. (ed.): Tertiary avian Freudenthal, M. ( 1971 ): Neogene vertebrates from the localities of Europe. Acta Univ. Carol. (Geologica) Gargano Peninsula, Italy. Scripta geol. 14: 1-19. 39: 529-533. Freudenthal, M. (1972): Deinogalerix koenigswaldi M1Novsk~, J. (1996b): Tertiary avian faunas of Eu- nov. gen., nov. spec., a giant insectivore from the rope. In: Mlfkovsk3), J. (ed.): Tertiary avian locali- Neogene of Italy. Seripta geol. 14: 1-19. ties of Europe. Acta Univ. Carol. (Geologica) 39: Freudenthal, M. (1976): Rodent stratigraphy of some 777-818. Miocene fissure fillings in Gargano. Scfipta geol. Ml~ovsl@, J. (in press): Early Miocene birds of Jebel 37: 1-23. Zelten, Libya. Acta Univ. Carol. (Geologica). Gaillard, C, (1908): Les oiseaux des Phosphorites du Mlfkovsk~, J., BNka, L. & Zemek, K. (1985): Mor- Quercy. Ann. Univ. Lyon (n.s,) 23: 1-178. phogenesis and the problem of morphospecies. In: Gaillard, C. (1939): Contribution h l'6tude des oi- M1Novsk¢, J. & Novfik, V.J.A. (eds.): Evolution seaux fossiles. Nouv. Arch. Mus. Lyon 15 (2): and morphogenesis: 201-211. Praha. 1-100. Mll'kovsk~,, J. & Hesse, A. (1996): Tertiary avian Haffer, J. (1995): Species versus lineages. In: Peters, localities of Germany. In: Mh'kovsk3), J. (ed.): Ter- D.S. (ed.): Acta palaeornithologica. Courier For- tiary avian localities of Europe. Acta Univ. Carol. schungsinst. Senckenberg 181: 303-309. (Geologica) 39: 619-647. Heissig, K. (1978): Fossilffihrende Spaltenftillungen Mourer-Chauvir6, C. (1987): Les Strigiformes Stiddeutschlands und die Okologie ihrer oligozg- (Ayes) des Phosphorites du Quercy (France): sy- hen Huftiere. Mitt. bayer. Staatssamml. Pal~iontol. st6matique, biostratigraphie et pal6obiogeogra- hist. Geol. 18: 237-288. phie. In: Mourer-Chauvir6, C. (ed.): L'6volution Hor~fiek, I. & Lo~ek, V. (1988): Palaeozoology and des oiseaux d'apr~s le tgmoignage des fossiles. the mid-European Quaternary past: scope of the Docum. Lab. G6ol. Lyon 99: 89-135. approach and selected results. Praha. Mourer-Chauvir6, C. (1994): A large owl from the J~nossy, D. (1991): Late Miocene bird remains from Palaeocene of France. Palaeontology 37: 339-348. Polgfirdi (W-Hungary). Aquila 98: 13-35. Mourer-Chauvir6, C. (1995): Dynamics of the avifau- Lambrecht, K. (1921): Pars 12. Ayes. In: Diener, C. na during the Paleogene and the early Neogene of (ed.): Fossilium catalogus. I. Animalia. Berlin. France. Settling of the recent fauna. Acta zool. Lambrecht, K. (1933): Handbuch der Palaeornitholo- cracov. 38: 325-342. gie. Berlin. Mourer-Chauvir6, C. (1996): Paleogene avian locali- Lydekker, R. (1891): Catalogue of the fossil birds in ties of France. In: M1Novsk~, J. (ed.): Tertiary the British Museum (Natural History). London. avian localities of Europe. Acta Univ. Carol. 39: Lydekker, R. (1893): On some bird remains from the 567-598. Miocene of La Grive-Saint-Alban, Department of Mourer-Chauvir6, C. & Sanchez Marco, A. (1988): Isere, France. Proe. zool. Soc. London 1893: 517- Prdsence de Tfto balearica (Ayes, Strigiformes) 522. dans des gisements continentaux du Pliocene de Mein, P. (1990): Updating of MN zones. In: Lindsay, France et d'Espagne. G6obios 21: 639-644. E.H., Fahlbusch, V. & Mein, P. (eds.): European Mourer-Chauvir6, C., J.A. Alcover, S. Moya & J. Neogene mammal chronology: 73-90. New York. Pons (1980): Une nouvelle forme insulaire d'ef- Miller, L. (1956): A collection of bird remains from fraie g6ante, Tyto balearica n. sp., (Ayes, Strigifor- the Pliocene of San Diego, California. Proc. Cali- rues), du Plio-Pleistoc~ne des Baleares. G6obios fornia Acad. Sci. 28: 615-621. 13: 803-811. Milne-Edwards, A. (1863): M6moire snr la distribu- Mourer-Chanvir6, C., Bour, R., Moutou, F. & Ribes, tion g6ologique des oiseaux fossiles et description S. (1994): Mascarenotus nov. gen. (Ayes, Strigi- de quelques esp6ees nouvelles. Ann. Sci. nat. (4) formes), genre end6mique 6teint des Mascareignes 20: 132-176. et M. grucheti n. sp., esp~ce 6teinte de La R6union. Milne-Edwards, A. (1869): Recherches anatomiques C. R. Acad. Sci. Paris (II) 318: 1699-1706. et pal6ontologiques pour servir ~ l'histoire des Newton, E. & Gadow, H. (1893): On additional bones oiseaux fossiles de la France. Vol. 2. Paris: Victor of the dodo and other extinct birds of Mauritius Masson et Fils. (+ Atlas 1869-1871). obtained by Mr. Th6odore Sauzier. Trans. Zool. Milne-Edwards, A. (1892): Sur les oiseanx fossiles Soc. London 13: 281-302. des d6pots ~ocenes de phosphate de chaux du Sud Olson, S.L. & Hilgartner, W.B. (1982): Fossil and de la France. C.R. 2 e Congr. ornithol, internat. 2: subfossil birds from the Bahamas. In: Olson, S.L. 60-80. (ed.): Fossil vertebrates from the Bahamas. Smith- Ml~ovsk~, J. ( 1992): The present state of knowledge son. Contrib. Paleobiol. 48: 22-60. of the Tertiary birds of Central Europe. In: Camp- Paris, P. (1912): Oiseaux fossiles de France. Rev. bell, K.E. (ed.): Papers in avian paleontology ho- frang. Omithol. 37: 283-298. noring Pierce Brodkorb, Nat. Hist. Mus. Los An- Peters, D.S. (1992): A new species of owl (Ayes: geles Co., Sci. Ser. 36: 433-458. Strigiformes) from the middle Eocene Messel oil J. Mlfkovsk~ • A new barn owl from the early Miocene 261

shale. In: Campbell, K.E. (ed.): Papers in avian Bubo incertus Milne-Edwards, 1892: 33. paleontology honoring Pierce Brodkorb. Nat. Hist. Nocturnavis incerta (Milne-Edwards): Mourer- Mus. Los Angeles Co., Sci. Ser. 36: 161-169. Chauvir6 1987:108 (new combination). Pomerol, C. & Premoli-Silva, I. (eds., 1986): Termi- Selenornis henrici (Milne-Edwards) [status uncer- nal Eocene events. Amsterdam. Rrmy, J.A., Crochet, J.Y., Sig6, B., Sudre, J., de tain] Bonis, L., Vianey-Liaud, M., Godinot, M., Harten- Otus Henrici Milne-Edwards, 1892: 63. berger, J.L., Lange-Badrr, B. & Comte, B. (1987): Asio Henrici (Milne-Edwards): Gaillard 1908:36 Biochronologie des Phosphorites du Quercy: Mise (new combination). 5jour des listes faunisfiques et nouveaux gisements Asio henrici (Milne-Edwards): Paris 1912:287 de mammif~res fossiles. In: Schmidt-Kittler, N. (spelling emended) (ed.): International symposium on mammalian Selenornis henrici (Milne-Edwards): Mourer- stratigraphy and paleoecology of the European Pa- Chauvir6 1987:113 (new combination). leogene. Mtinchner geowiss. Abh. (A) 10: 169- Necrobyas harpax Milne-Edwards 188. Necrobyas harpax Milne-Edwards, 1892:61. Rich, P.V. & Bohaska, D. (1976): The world's oldest Necrobyas Rossignoli Milne-Edwards, 1892:63 owl: a new strigiform from the Paleocene of south- (here synonymized). western Colorado. In: Olson, S.L. (ed.): Collected Necrobyas rossignoli Milne-Edwards: Paris 1912: papers in avian paleontology honoring the 90th 287 (spelling emended) birthsday of Alexander Wetmore, Smithson. Con- trib. Paleobiol. 27: 87-93. Necrobyas arvemensis (Milne-Edwards) Rich, P.V. & Bohaska, D. (1981): The Ogygoptyngi- Bubo arvernensis Milne-Edwards, 1863: 458. dae, a new family of owls from the Paleocene of Necrobyas Edwardsi Gaillard, 1939:8 (here syn- North America. Alcberinga 5: 95-102. onymized). Rothschild, W. (1907): Extinct birds. London. Paratyto arvernensis (Milne-Edwards): Brodkorb Salotti, M., Bailon, S., Bonifay, M.-F., Courtois, J.- 1970:159 (new combination). Y., Dubois, J.-N., Ferrandini, J., Ferrandini, M., La Necrobyas edwardsi GaiUard: Brodkorb 1971: 220 Milza, L.-C., Mourer-Chanvirr, C., Popelard, J.- (spelling emended) B., Quinif, Y., Rral-Testud, A.-M., Miniconi, C., Necrobyas arvernensis (Milne-Edwards): Mourer- Pereira, E. & Persiani, C. (1997): Castiglione 3, un Chauvir6 1987:91 (new combination). nouveau remplissage fossilifrre d'~ge P16istoc~ne moyen dans le karst de la r6gion d'Oletta (Haute- Prosybris media (Mourer-Chauvirr) Corse). C.R. Acad. Sci. Paris (IIa) 324: 67-74. Necrobyas medius Mourer-Chauvirr, 1987: 104. Schmidt-Kittler, N. (1987): European reference le- Prosybris media (Mourer-Chauvirr): Ml~ovsk~, vels and correlation tables. In: Schmidt-Kittler, N. this paper (new combination). (ed.): International symposium on mammalian Prosybris antiqua (Milne-Edwards) stratigraphy and paleoecology of the European Pa- Strix antiqua Milne-Edwards, 1869: 498. leogene. Mtinchner geowiss. Abh. (A) I 0:13-19. Prosybris antiqua (Milne-Edwards): Brodkorb Schneider, W. (1977): Schleiereulen (Tytonidae). 1970:159 (new combination). Wittenberg Lutherstadt. Necrobyas minimus Mourer-Chauvirr, 1987:105 Wetmore, A. (1920): Five new species of birds from (here synonymized) cave deposits in Porto Rico. Proc. biol. Soc. Was- hington 33: 77-82. Tyro sanctialbani Lydekker Wetmore, A. (1922): Remains of birds from caves in Strix sancti-albani Lydekker, 1893: 518. the Republic of Haiti. Smithson. misc. Coll. 74 (4): Strix sancti albani Lydekker: Ennouchi 1930:66 1-4. (spelling emended). Wetmore, A. (1937): Bird remains from cave deposits Strix Sancti-albani Lydekker: Gaillard 1939:82 on Great Exuma Island in the Bahamas. Bull. Mus. (spelling emended) comp. Zool. 80: 427-441. Tyto sanctialbani (Lydekker): Ballmann 1969b: Wolters, H.E. (1975-1982): Die Vogelarten der Erde. 191 (new combination and spelling emended). Hamburg. Tyro campiterrae J~nossy, 1991:25 (here synonymized). Accepted: 03 February 1998 Tyro balearica Mourer-Chanvir6 et al. Tyro balearica Mourer-Chauvir6, Alcover, Moya Appendix. Systematic list of & Pons, 1980: 804. Tertiary barn owls Tyto gigantea Ballmann (subfamily Tytoninae) Tyto gigantea Ballmann, 1973: 37. Tyto robusta Ballmann, 1973:33 (here synonyrrfiz- Nocturnavis incerta (Milne-Edwards) ed).