In this chapter, I discuss the Neogene and Early Pleistocene fossil record of Coturnicini and provide a synthesis of the evolutionary history of the group in Eurasia based on paleontological data. I discuss the early stages of the evolution of the group, paying special attention to Tologuica, the well-known early representatives of Coturnicini from Mongolia and Eastern Siberia. The coexistence of the genera Chauvireria and Plioperdix in the Late Miocene and Early Pliocene of Eastern Europe mark a separate well-represented stage in the evolution of the group in Eurasia. Possible distinctive specialization and different adaptations of these taxa are briefly discussed. In Western and Central Europe, coturnicines are represented by the separate genera Palaeocryptonyx and Palaeoperdix, both known starting from the Middle Miocene. The genus Palaeoperdix comprises quail-like forms, whereas the species of Palaeocryptonyx are phylogenetically more distant from Coturnix and relatives. The Late Pliocene species Chauvireria minor and Palaeocryptonyx capeki are considered as the most specialized taxa within these genera (both probably confined to grassy vegetation landscapes), and the Afro-Iberian taxon Marocortyx is considered a transitional form (showing similarities with modern Margaroperdix) between Alectoris-like coturnicines and highly-specialized quails. Africa may have played an important role in the evolution of the group, although some of the diversification of Coturnicini obviously occurred in Asia. The oldest occurrences of modern taxa are discussed, with a special emphasis on the evolution of Coturnix quails, which first occur in Eurasia in the Late Pliocene of Mongolia, but spread across Europe only in the Calabrian of the Early Pleistocene.
The Cuban macaw Ara tricolor (Bechstein, 1811) is an extinct species of large parrots. Its historical distribution and ecology are poorly understood. To date, only three late Quaternary paleontological and one archeozoological (17th–18th centuries) finds of the species have been described from central Cuba. A new (fourth) fossil find of the Cuban macaw is described and is a fragmentary carpometacarpus from Upper Pleistocene layers of the El Abrón Cave in the Pinar del Río province. The find provides the first documented evidence of the species from the western part of Cuba. The associated fauna indicates that the Cuban macaw lived in mosaic, semi-open marshy landscapes, which were probably similar to those in the vicinity of Ciénaga de Zapata in modern times.
Old World vultures (Aves: Accipitridae: Gypini) are large diurnal birds of prey, the characteristic inhabitants of open biotopes in Africa and southern Eurasia. Fossil remains of vultures are rather rare; the evolution of the group is poorly studied. This article describes the skull and tarsometatarsus of the large vulture Torgos platycephalus sp. nov. from the Upper Pleistocene of the Binagadi locality (Azerbaijan). This is the first fossil record of the genus in the Caucasus and second confirmed record outside its current range (Torgos sp. is also known from the Middle Pleistocene of China). The coexistence of three species of large scavengers in the Late Pleistocene of the Absheron Peninsula (in addition to Torgos platycephalus, Aegypius monachus and Gyps fulvus are also known from the Binagadi locality) can be explained by the richness of the food supply, which included a noticeable diversity of large mammals.
The Great Lakes Depression in northwestern Mongolia contains widespread Late Neogene continental deposits, reaching thicknesses of several hundred metres. The Late Neogene Khyargas Nuur formation (also known as the Khirgis-Nur formation) has a significant biostratigraphic value for Central Asia, as it encompasses a unique palaeontological record and has a wide geographical distribution in the region. The stratotype of this formation was described in the Khyargas Nuur 2 (=Khirgis-Nur-2) section on the northern shore of Khyargas Lake. Bio- and magnetostratigraphic studies conducted in the 1970s and 1980s identified the Miocene-Pliocene boundary (corresponding to the Turolian and Ruscinian biochrons) within Unit A, or the Lower Khyargas Nuur subformation. This study aims to refine the age and stratigraphic subdivisions of the Khyargas Nuur formation in light of new data. Examination of the geology of the Lower Khyargas Nuur Subformation stratotype, along with a revision of the associated mammalian assemblage (carnivores, ungulates, and rodents), supports the conclusion that this faunal complex corresponds to the Late Turolian (MN13) of the continental biochronological scale, thus assigning this section to the terminal Miocene. These findings provide new insights into the geological history of the Great Lakes Depression and the faunal history of Central Asia.
Buttonquails (family Turnicidae of the order Charadriiformes) are a morphologically specialized group of small, predominantly tropical birds of open landscapes, which is extremely poorly represented in the fossil record. The article describes a fragmentary humerus of a buttonquail from the Lower Pleistocene of the Taurida Cave in central Crimea. This is the first find of the family Turnicidae in Eurasia in a chronological interval from the Pliocene through the Middle Pleistocene. The find highlights the limited nature of available information on the taxonomic composition of Early Quaternary Eurasian avifaunas, even at the family level, and sheds light on the Late Cenozoic evolutionary history of Turnicidae.
The introduction provides general information on the diversity, evolutionary history, and fossil record of quails and allies (Aves: Phasianidae: Coturnicini). Estimates on the dates of divergence of particular taxa of Coturnicini based on molecular data are critically reviewed. It is concluded that most of the divergence dates are too old, since they are based on inappropriate fossil calibrations. When appropriate calibrations are used, estimated divergence dates are much more consistent with the overall fossil record of this group.
Remains of Lyrurus partium Kretzoi, 1962 and “Lagopus” atavus Jánossy, 1974 (tribe Tetraonini, family Phasianidae, order Galliformes) are described from the Lower Pleistocene of the Taurida Cave (Crimea); these are the oldest finds of the grouse in Eurasia outside of Central and Eastern Europe (Bulgaria, Hungary, and Poland). The discovery of the primitive black grouse Lyrurus partium indicates that these birds were part of the savanna-like faunal associations of the Northern Black Sea region (along with Otididae and Struthionidae) in the Early Pleistocene. The origin and evolutionary history of the modern Caucasian black grouse T. mlokosiewiczi Taczanowski, 1875 is discussed. The complete tarsometatarsus of “Lagopus” atavus from the Taurida Cave indicates a separate generic status of this poorly studied Plio-Pleistocene form, which is here transferred to the genus Paralyra gen. nov. Therefore, the presence of ptarmigans (genus Lagopus) in the preglacial Plio-Pleistocene faunas of Eastern Europe is not confirmed.
This chapter provides a short account of the comparative osteology of modern members of Coturnicini. Coracoids, humeri, carpometacarpi, and tarsometatarsi are used in this study as the most diagnostic skeletal elements and those which are often available as fossils. Main diagnostic characters are indicated for these bones. The osteological nomenclature is provided. Additionally, the phylogenetic relationships between several fossil taxa of Phasianidae are investigated. Preliminary results of a new phylogenetic analysis for Galliformes are given. The fossil taxa Tologuica, Chauvireria, and Plioperdix are recovered as close relatives of modern Coturnix quails. The fossil Palaeocryptonyx is ambiguously placed as a basal taxon within Coturnicini.
The article describes the first find of a bird from the Paleogene of Siberia. A fragment of tibiotarsus from the Eocene Tavda Formation of the Tyumen Region (Western Siberia) is assigned to Procellariiformes. The bird is morphologically closer to Procellariidae, but comparable in size to albatrosses (Diomedeidae) and is assumed to represent the stem members of the family. The find indicated for the first time that either stem albatrosses or similar large Procellariiformes could have had a worldwide distribution as early as the Eocene.
Material on ducks from the Lower Pleistocene (about 1.8 Ma) of Taurida Cave in the Crimea represents forms new to the locality: Melanitta kirbori sp. nov., Marmaronetta angustirostris, Spatula querquedula, Anas aff. A. acuta, and Anas aff. A. platyrhynchos. In addition, new material on the previously known taxa Spatula praeclypeata Zelenkov, 2022 and Tadorna petrina Kurochkin, 1985 is described. The assemblage of ducks from Taurida Cave is the oldest representative fauna of Anatidae for the Early Pleistocene of Europe. The finds of the genera Melanitta and Marmaronetta are the oldest in the fossil record on a global scale. The finds of ducks from the groups of mallards (Anas aff. A. platyrhynchos) and pintails (Anas aff. A. acuta) are the oldest known for these lineages, marking the dispersal of the ancestors of modern Palearctic species from Africa to Eurasia. The evolutionary history of the anatid taxa found in the cave is discussed.
This paper describes remains of fossil gray partridges from the Lower Pleistocene (1.8–1.5 Ma) deposits of the Taurida Cave in Central Crimea. The smaller gray partridge is referred to the fossil species Perdix jurcsaki Kretzoi, 1962. Another phasianid bird from the cave, which is partly similar to modern Perdix but larger than the latter, is described as Enkuria voinstvenskyi gen. et sp. nov. The spatiotemporal stability of the Early–Middle Pleistocene European gray partridge morphotype makes it possible to synonymize the fossil species P. jurcsaki and P. palaeoperdix Mourer-Chauviré, 1975. Earlier fossil finds of the genus Perdix from the Upper Pliocene–Lower Pleistocene of Eastern Europe and Asia are revised. Perdix margaritae Kurochkin, 1985 from the Upper Pliocene of Mongolia and Transbaikalia is considered a junior synonym of Lophura inferna Kurochkin, 1985. The oldest confirmed find of P. jurcsaki comes from the lower Gelasian (about 2.4 Ma) of the Northern Black Sea region (Kryzhanovka-2 locality; Ukraine). A dispersal of gray partridge to the west of Europe in the second half of the Early Pleistocene is shown. The discovery of the genus Enkuria confirms the existence of the fossil diversity of Perdicini s.s. in the Plio-Pleistocene of Northern Eurasia. Phasianus etuliensis Bocheński et Kurochkin, 1987 from the Lower Pliocene of Moldova is also here assigned to this genus. In addition, the validity of the fossil Early Pleistocene phasianids Francolinus subfrancolinus Jánossy, 1976 and Alectoris baryosefi Tchernov, 1980 is confirmed.
This chapter provides a systematic description of all confirmed or reliable finds of fossil coturnicines (Aves: Galliformes: Phasianidae: Coturnicini) from the Miocene to the Lower Pleistocene of Eurasia, as well as North Africa (Morocco). Previously published data are reviewed and newly described materials come from the Miocene to the Lower Pleistocene of Eastern Europe (Ukraine and European part of Russia) and Eastern Siberia (Baikal), as well as from the Upper Pliocene of Mongolia. The oldest member of the Coturnicini clade (Tologuica vetusta sp. nov.) is described from the uppermost Lower Miocene-lowermost Middle Miocene of Baikal area (Tagay locality). Members of the fossil genus Tologuica are present in the upper Middle Miocene of Mongolia and the fossil genus Palaeoperdix, which is here revised and transferred to Coturnicini, is known from the upper Middle Miocene of Europe (Palaeoperdix longipes Milne-Edwards, 1869 and Palaeoperdix grivensis (Ennouchi, 1930) comb. nov.). Plioperdix (“Palaeocryptonyx”) hungarica (Jánossý, 1991) from the Upper Miocene of Hungary is transferred to Palaeoperdix. Mioryaba magyarica Zelenkov, 2016 is shown to be similar to modern Perdicula. Several Upper Miocene through Lower Pleistocene coturnicine taxa are assigned to the fossil genera Chauvireria and Plioperdix, with several new species erected: Chauvireria egorovkensis sp. nov., Chauvireria axaina sp. nov., and Plioperdix boevi sp. nov. (all of them from the Upper Miocene of North Black Sea area). The Late Pliocene–Early Pleistocene taxon Plioperdix (“Francolinus”) capeki (Lambrecht, 1933) is transferred to Palaeocryptonyx, and the Late Pliocene taxa Palaeocryptonyx novaki Sánchez Marco, 2009 from Spain and Plioperdix africana Mourer-Chauriré Geraads, 2010 from Morocco are transferred to a new genus, Marocortyx gen. nov., which is considered a morphologically transitional form between Alectoris and Margaroperdix. The new finds of Palaeocryptonyx donnezani Depéret, 1892 are described, which were previously partly treated as Plioperdix pontica. Some previously published finds of P. donnezani are revised. The oldest find of Coturnix quails in the region is from the Upper Pliocene of Mongolia (Shaamar) and is described as Coturnix augustus sp. nov. Coturnix coturnix is further described from the Lower Pleistocene of Crimea (Taurida Cave).
A revision of small ducks (the size of the modern teal Anas crecca or smaller) from the middle Miocene of France (Sansan locality) and Mongolia (Sharga locality) clarified the taxonomic status and systematic position of the well-known species Anas velox Milne-Edwards, 1868 and Anas soporata Kurochkin, 1976. It is shown that three small members of the family Anatidae are present in the fauna of the Sansan locality: Anas velox is a diving duck, partly similar to modern Histrionicus, but smaller–here this species is transferred to the fossil genus Protomelanitta Zelenkov, 2011 (basal Mergini). A somewhat smaller taxon from Sansan belongs to the ecological group of dabbling ducks, and is identified as Anas soporata, a species that was previously described from Mongolia and here transferred to the genus Mioquerquedula Zelenkov et Kurochkin, 2012. In addition, yet another very small duck of unclear systematic position is present in the fauna of Sansan. New materials on Mioquerquedula soporata comb. nov. and M. minutissima Zelenkov et Kurochkin, 2012 are also described from the middle Miocene of Mongolia.
Central Asia is a region that, owing to its geographical position, has always been at the intersection of cultures. In the Fergana Valley, located in the very heart of Central Asia, the Late Pleistocene–Early Holocene is characterized by the expansion and intensification of human settlements. This cultural event has previously been linked to global climate changes that took place during the Pleistocene–Holocene transition. In order to reconstruct local climatic conditions in the area during this period, it is worth studying the fossil vertebrate faunas from late Quaternary sites in the Fergana Valley. The Obishir-5 rock shelter, dating to the Late Pleistocene–Early Holocene period, discovered in the middle of the 20th century, is a key archeological site in the valley. To date no thorough zooarcheological research has been done on the excavated faunal remains. Here we present the results of the first faunal analysis of the small vertebrates found at this site. We studied the taxonomic composition of remains, which was then used to make paleoenvironmental reconstructions. In total, we identified 13 small mammal species and three bird species. In addition, the assemblage contained one amphibian and two reptile taxa. The faunal composition suggests that during the final Pleistocene and Early Holocene the study area enjoyed a stable climate with vegetation dominated by open biotopes (alpine meadows, steppes, semi-deserts). Its location makes the Fergana Valley an example of a geographically isolated area for a terrestrial fauna which can exist for a long time without being influenced by biotic factors from neighbouring territories.
Sandgrouse (Pteroclidae, Pterocliformes) are specialized ground birds of open arid landscapes with a very poorly studied evolutionary history. In the late Pliocene–early Pleistocene, Pteroclidae are known only from few localities in Southern Europe. The article describes a relatively large fossil sandgrouse from the early Pleistocene of the Taurida cave in the Crimea. This is the first record of Pteroclidae in the ancient faunas of the Black Sea region and Eastern Europe. The unusual structure of the tibiotarsus makes it possible to describe the fossil form from Taurida Cave as a new species, Pterocles bosporanus sp. nov.
Evolutionarily advanced waterfowl (the modern family Anatidae and its stem members in the order Anseriformes) are a successful bird group, well represented in faunas globally since the late Oligocene. However, the pre-late Oligocene history of the evolutionary lineage of Anatidae remains largely unexplored, as these birds are very rare in earlier faunas. This paper describes a remarkable diversity of waterfowl from the lower Oligocene of Kazakhstan (Central Asia), which includes two members of Anatidae, as well as stem-anatids referred to Romainvilliidae and Paranyrocidae. The latter family is represented by the enigmatic swan-sized Cygnopterus, the genus revised here, with new materials confirming its ordinal and familiar assignment. Anatidae are represented by the globally oldest diagnosable taxa for this family, including species in the fossil genus Mionetta, previously known from upper Oligocene-Middle Miocene strata. The paper further addresses two upper Eocene anseriform taxa from Kazakhstan. These data show that diverse assemblages of advanced anseriforms existed in Asia as early as early Oligocene. The presence of several stem-anatid taxa in the late Eocene faunas of Central Asia supports the hypothesis of a possible Eurasian (or Northern Continents) origin of Anatidae, although the subsequent diversification of the group could have occurred in the Australian region.
Taxonomic diversity of Late Cretaceous hesperornithids (Aves: Hesperornithidae) of European Russia and Eastern Europe as a whole remain poorly understood, and the morphology of these large flightless birds is poorly known. New finds of Hesperornithidae in the Karyakino locality (Saratov oblast, Russia) confirm the coexistence of two forms of these flightless seabirds in the Campanian (mid-Late Cretaceous) of the Lower Volga region. A femur is described for the first time for Hesperornis rossicus Nessov et Yarkov, 1993, indicating that this large species is morphologically distinct from North American H. regalis Marsh, 1872.
The results of a taxonomic and morphological revision of Mionetta natator (Milne-Edwards, 1867) from the lower Miocene of Western and Central Europe are presented. It is shown that the collections of small ducks from the Saint-Gérand-le-Puy localities (France) do indeed contain a small species of the genus Mionetta, which is here described as Mionetta defossa sp. nov. However, Mionetta natator represents a separate taxon, and is here assigned to the genus Caerulonettion gen. nov., showing morphological similarities with extant Malacorhynchus and fossil Mioquerquedula spp. The first small ducks from the lower Miocene of Kazakhstan are also described (Aral Formation, the Agyspe and Altynshokysu localities; Akzhar Formation, the Golubye Peski locality), tentatively attributed to Mionetta defossa and Mionetta sp. The diversity and relationships of small ducks of the Early and Middle Miocene of Eurasia are discussed.
Remains of small ducks are described from the boundary Early–Middle Miocene deposits of Tagay (Baikal Region), the only locality in Asia with a representative bird fauna from the Miocene climatic optimum. The new taxa Mioquerquedula palaeotagaica sp. nov. and Tagayanetta palaeobaikalensis gen. et sp. nov., which correspond to modern Anas crecca in size, as well as an even smaller duck, Selenonetta lacustrina gen. et sp. nov., are described. The genus Mioquerquedula is revised; “Anas” integra Miller, 1944 from the Lower Miocene of North America is transferred to this genus. Tagayanetta gen. nov. is considered here as an evolutionarily more advanced genus than Mioquerquedula and is probably close to Anatini. Selenonetta gen. nov. is considered as a taxon close to the divergence between Mergini and other Anatinae. A similar form (possibly the same species) is present in the Sansan locality (France).
The oldest remains of anseriform birds for the Early Pleistocene of the Northern Black Sea region are described based on materials from the Taurida Cave (central part of the Crimean Peninsula, 1.9–1.5 Ma). A fragmentary skeleton of a shelduck (Tadornini) is attributed to the fossil stone shelduck Tadorna petrina Kurochkin, 1985, originally described from the Upper Pliocene of Transbaikalia and morphologically close to the modern Ruddy Shelduck T. ferruginea. This is the oldest find of the Ruddy Shelduck lineage in Europe, which sheds light on the paleobiogeographical distribution and evolution of these shelducks in the Late Cenozoic. Tadorna petrina is considered as the most likely stem representative of the clade, which includes the modern species T. ferruginea, T. cana, T. tadornoides, and T. variegata. The new fossil Shoveler Spatula praeclypeata sp. nov. from the Taurida Cave is the oldest find of this lineage in the fossil record and considered as a stem taxon of the living species S. smithii, S. rhynchotis, and S. clypeata. The faunal association of Ruddy Shelduck and Shoveler is characteristic of many Middle and Late Pleistocene avifaunas of Europe; Crimean finds confirm its more ancient and, probably, eastern origin.