Ornis Hungarica 2014. 22(2): 65–125. DOI: 10.2478/orhu-2014-0019 Fossil and subfossil bird remains and faunas from the Carpathian Basin Jenő (eugen) Kessler Jenő (Eugen) Kessler 2014. Fossil and subfossil bird remains and faunas from the Carpathian Basin. – Ornis Hungarica 22(2): 65–125. Abstract The author summarizes the knowledge of fossil and subfossil bird life from the Car- pathian Basin, of all geological ages, site by site. After a historical overview, he presents the Mesozoic, Tertiary and Quaternary bird fauna, based on a holistic reference material consisting of 196 titles indi- cated in the bibliography, including papers in English (64), Hungarian (50), German (46), Romanian (26), Cro- atian (9) and Polish (1) languages. The text is supplemented with maps of fossiliferous sites from different ages and a list from 341 paleontological and archaeological sites on species of the Carpathian Basin, respectively. The number of taxa reaches 845, including 189 extinct taxa (two orders, four families, nine genera – five ichnotaxa of which (154 species, five ichnotaxa and 10 subspecies) were described from the Carpathian Basin, primarily. Most significant records include the Mesozoic taxa (Eurolimnornis, Palaeocursornis, Elopteryx), the new Neo- gene songbirds species and the presence of predecessors of recent European grouses, Bustards and Corvidae spe- cies from the Pliocene and Early Pleistocene in the Carpathian Basin. Keywords: Carpathian Basin, paleo-ornithological studies, fossil sites, evolution Összefoglalás A szerző összefoglalja a Kárpát-medence fosszilis és szubfosszilis madárvilágáról szóló ismere- teket, mind koronként, mind lelőhelyenként. A történelmi visszatekintés után ismerteti a mezozoikumi, a har- mad- és negyedidőszaki lelőhelyek madárfaunáját egy teljességre törekvő, angol (64), magyar (50), német (46), román (26), horvát (9) és lengyel (1) nyelvű, 196 tételből álló hivatkozáslista alapján. A kréta és harmadidősza- ki, a kora- és középső-pleisztocén, a késő-pleisztocén, valamint a késő-glaciális és holocén kori lelőhelyek tér- képei, valamint (mellékletként) a meghatározott taxonok rendszertani és időrendbeli táblázata egészíti ki a tanul- mány szövegét. A Kárpát-medencéből 341 őslénytani és régészeti lelőhelyről ismerünk fosszilis és szubfosszilis madármaradványokat. A taxonok száma eléri a 845-öt. Ebből 189 kihalt taxon (két rend, négy család, kilenc ge- nus – ebből öt ichnotaxon (154 faj, öt ichnotaxonnal és 10 alfaj) először a Kárpát-medencéből került leírásra. Legjelentősebb leletek a mezozoikumiak (Eurolimnornis, Palaeocursornis, Elopteryx), de igen fontos a neogen kori énekesek és a recens európai fajdok, túzokok és varjúfélék fosszilis elődjei jelenlétének kimutatása is a Kár- pát-medence pliocénjében és kora-pleisztocénjében. Kulcsszavak: Kárpát-medence, madárőslénytani kutatások, madárfossziliák, evolúció, fosszília-lelőhelyek Department of Paleontology, Eötvös Loránd University, 1117 Budapest, Pázmány Péter sétány 1/c, Hungary, e-mail: [email protected] Introduction vided adequate conditions for bird life, pre- sumably, in the time when they appeared on The paleogeographic conditions in the Car- the Earth. The area was covered by more or pathian Basin’s ancient area (the tectonic less widespread masses of water (lakes, in- plates which constitute the actual territory land and open seas) from the Triassic to the were not necessarily on their present-day Neogene period, but sometimes it had vast place all the time) are believed to have pro- littoral zones with lagoons and archipelagos 66 ORNIS HUNGARICA 2014. 22(2) as well. We have a lot of evidence of conti- In this region, another characteristic of that nental life already from the Triassic for both 200 million years long period between the the plant and animal kingdoms. There are Triassic and the Quaternary was a warmer well-known plant marks and relics develo- climate. This can be seen from the compo- ped in the Mecsek and Királyerdő Moun- sition of both the flora and vertebrate fauna. tains (Munţii Pădurea Craiului) from the For example, several vertebrates, which to- Early Jurassic. However, animal remains day are characteristic to the tropics (primeval such as footprints of dinosaurs in Mecsek, reptiles, crocodiles, turtles, birds and even nothosaurs described in the Báród Basin, mammals), lived here. The diverse and lush a sea-protocrocodile skeleton in Gerecse, vegetation with its insects, as well as the fau- Early Cretaceous remains of vertebrates of na of the vast aquatic surroundings, provided Cornet in Királyerdő, fossils of dinosaurs, rich food sources for many, including birds, crocodiles, lizards but even Mammalia from allowing them to exist. Their presence is indi- the Late Cretaceous in Erdély (Transylva- cated by findings already from the Mesozoic, nia) and in the Bakony. As a consequence of but we have plentiful fossils from the Tertiary paleogeographic conditions, terrestrial life and especially from the Neogene. As a conse- forms are poorly represented (e.g. there are quence of the extensive karstification, in nu- fewer bird fossils). merous caves, rich bird fossils from the Qua- This is plausible because the Carpathian ternary were discovered. These fossils allow Basin did not even exist at that time. It was us to easily trace the changes of the avifauna represented by islands or microcontinents during the succeeding glacial and interglacial located on three tectonic plates between Eu- periods, up to the present situation. rope and Africa. Plant fossils indicate that We discuss in this paper the evolution of climate was warmer than in the rest of Eu- this avifauna based on the available litera- rope, but their area has limited both the va- ture and the bird fossils examined and de- riety of forms of life and the possibilities of termined by us. fossilization. From the Neogene and espe- cially after the formation of the basin in the List of collections: present-day location, during and following GIB = Geological Institute Bucureşti, Ro- the uprising of the ring of mountains and the mania formation of the internal sea which later be- IPUW = Institute of Paleontology, Universi- came a lake, in many places, there appeared ty of Wien, Austria adequate conditions for fossilization. As a EMNH = Eötvös Museum of Natural His- consequence, in many places, even bones of tory, Eötvös Loránd University, Budapest, birds could be found. From the beginning Hungary of the Quaternary, the life forms in the Car- LPUB = Laboratory of Paleontology, Uni- pathian Basin became more and more simi- versity Bucureṣti, Romania lar to the present-day ones. This flora and MÁFI = Magyar Állami Földtani Intézet fauna had many similarities to those de- (Geological Institute of Hungary, new scribed in other parts of Europe but also had Geological and Geophysical Institute of a lot of endemic species. From the Mesozo- Hungary: GGIH, Budapest, Hungary) ic, even the occurrence of dwarfism, a cha- MEMEK = Museum ‘Erdélyi Múzeum racteristic of archipelagos, is demonstrated. Egylet’ Kolozsvár, Romania J. Kessler 67 MMB = Moravian Museum Brno, Czech fying the findings from the Nándori Cave Republic (Téglás 1880). With paleontological re- MMP = Museum Municipal Pásztó, Hun- search making rapid headway in the late gary ninetieth-early twentieth century, the year MPUBBC = Museum of Paleontology Uni- 1904 was a significant one regarding avian versity ‘Babeṣ-Bolyai’ Cluj – Kolozsvár, paleontology, when Mihály Tóth discovered Romania the site on the southwest versant of Somló MTM = Magyar Természettudományi Hill near Oradea. Excavation of the remains Múzeum (Department of Paleontolo- of the cave system formerly known as gy and Geology of the Hungarian Natu- Püspökfürdő, then as Betfia, was initiated by ral History Museum, Budapest, Hungary Tivadar Kormos. He was also the one to – HNHM) publish the avian fossils’ preliminary classi- MṬCO = Museum ‘Ṭării Criṣurilor Oradea’, fication (Kormos 1911, 1913). In-depth clas- Nagyvárad, Romania sification was made by Waclav Čapek Czech NHML = Natural History Museum, Lon- osteologist (Čapek 1917). In 1905, Zoltán don, England Szilády gathered material (avian bones NHMW = Natural History Museum Wien, among others) from the Lucsia Cave near Austria Aranyos-Szohodol in Transylvania. These remains have only been found in 2000 in the The short history of avian paleontology collection of the Department of Geology and in the Carpathian Basin Paleontology of the Hungarian Natural His- tory Museum by Erika Gál and identified re- According to the earliest known data (Neu- cently (Gál 2002a). Bones found in the ca- geboren 1850), Michael Ackner collected vern of Dealul Şprenghi in Braṣov were songbird remains around the middle of the classified by Ferenc Toula (Toula 1909). nineteenth century from the Popláka chasm A few years later, Gyula Éhik studied the near Sibiu in Transylvania. The next finding, same site (Éhik 1913). In 1915, Ottokár an imprint of a feather, was reported from Kadić gathered material from Zoltán Cave in the Miocene of Oltszakadát and Braṣov in Băile Herculane, Romania. Findings of the 1855 (Andrae 1855). Hermann von Meyer latter two excavations mentioned were also described a finch (Fringilla radobojensis) classified by Erika Gál (Gál 2002a). Several from the Miocene of Radoboj, Croatia (von fossils collected in the meantime have been Meyer 1865). A few years later, Samu Róth transported to foreign museums. From the excavated caves I. and III. of Novi Hill in material purchased from Ferenc Nopcsa and Szepes