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A new species of eagle (Aves: Accipitridae) close to the Steppe Eagle, from the Pleistocene of Corsica and Sardinia, France and Italy
Abstract
New data about fossil eagles of the genus Aquilo, from Corsica and Sardinia are presented. They lead to the description of Aquila nipaloides n. sp., an extinct species of large size, related to the A. nipalensis-rapax-heliaca- adalberti clade. The finds represent the first evidence of a member of this clade from the fossil record. A complete mandibula particularly shows its affinity with A. nipaknsis. Some elements, like the femur, indicate specific apomorphies. This thus increases the endemism in the Pleistocene record of these islands. It also indicates the presence of A. nipalensis, or a close ancestor, in Europe during at least part of the Pleistocene. This is well in accordance with a widespread steppe-tundra environment. This fact also indicates that it will be necessary to take this species into account when identifying fossils from this period in Europe. It is emphasized again that A. rapax is much distinct from A. nipalensis. We here synonymize Buthierax pouliani KRETZOI, 1977 (Middle Pleistocene, Petralona, Greece) with Buteo rufinus jansoni MOURER-CHAUVIRÉ, 1975b, and we consider A. chrysaetos simurgh WEESIE, 1988 (Upper Pleistocene, Crete, Greece) as Aves incertae sedis pending further investigations.
... 2: Aquila claudeguerini n. sp., measurements of the tarsometatarsus, in mm, compared to recent and extinct species of eagles. 1/ After Louchart et al. (2005), except for the width in the middle which is not given by these authors; 2/ After Ballmann (1973), some measurements have been taken directly on figure 2. Tab. 2 : Aquila claudeguerini n. sp., dimensions du tarsométatarse, en mm. 1/ D'après Louchart et al. (2005), excepté pour la largeur au milieu qui n'est pas indiquée par ces auteurs ; 2/ D'après Ballmann (1973), certaines mesures ont été prises directement sur la figure 2. ...
... 1/ After Louchart et al. (2005), except for the width in the middle which is not given by these authors; 2/ After Ballmann (1973), some measurements have been taken directly on figure 2. Tab. 2 : Aquila claudeguerini n. sp., dimensions du tarsométatarse, en mm. 1/ D'après Louchart et al. (2005), excepté pour la largeur au milieu qui n'est pas indiquée par ces auteurs ; 2/ D'après Ballmann (1973), certaines mesures ont été prises directement sur la figure 2. ...
... The tarsometatarsus is incomplete but, measuring the distance between the lateral proximal vascular foramen and the distal end, compared to the same distance in a recent specimen of A. chrysaetos, the total length can be estimated at 132 mm. In a recent population of 26 individuals, the maximal length of the tarsometatarsus is 112.1 mm (Louchart et al., 2005) (tab. 2). ...
The Ceyssaguet avifauna includes four taxa, among which a new species of the genus Aquila, dedicated to Claude Guérin, and characterized by a larger size and different proportions from the other species of this genus. It also includes the recent species Phalacrocorax carbo, the Great Cormorant, the presence of which can be explained by the proximity of a lake and also of the Loire River. The presence of Accipiter gentilis, the Northern Goshawk, indicates a wooded environment, which agrees well with the results of the pollen analysis.
... It lies over a thick calcitic floor, which levels at 1.32 m away from the N-S, W-E grid (Figs 6, 7) and slopes eastward (Fig. 5). U/Th dating of a stalagmite overlying the floor indicates that the overlying sediment is younger than 1 to 1.5 Ma. LOUCHART et al., 2005. In addition, remains of the large extinct insular shrew Episoriculus corsicanus BATE, 1944 (Soricidae) are extremely abundant. ...
... mandibular symphysis and pedal phalanges for A. nipaloides; complete tarsometatarsus and pedal phalanges for B. insularis), despite the fragmentary nature of the remains. Both species are known in Corsica and Sardinia from the Middle Pleistocene (Corsica only for the eagle) to the Late Pleistocene (for both islands) (LOUCHART 2002;LOUCHART et al. 2005). Within U.S. 4 to 22, B. insularis is represented by a minimum of nine individuals, and A. nipaloides by a minimum of two individuals (plus 10 Aquila sp. ...
... The two extinct species A. nipaloides and B. insularis are useful as chronological indicators. Hitherto, the most recent dated occurrences are as follows: A. nipaloides: latest Pleistocene of Corsica and Sardinia (Corsica: Castiglione 3PL; Sardinia: Corbeddu,Rapaci and Dragonara;LOUCHART 2002;LOUCHART et al. 2005); B. insularis: early Holocene (9500-8500 BPcal in Corsica; undated Holocene in Sardinia; ...
... H. arborea) and two reptiles (Coluber and Natrix) similar to the species living at present in Sardinia (Kotsakis, 1980). The avifauna is more diversified and includes 26 taxa, many of which are predators (Gavia sp., Phalacrocorax aristotelis, Ardea sp., Anser sp., Branta sp., Anatinae indet., Gypaetus barbatus, Aegypius monachus, Aquila nipaloides, Aquila chrysaetos, Aquila sp., Milvus milvus, Buteo buteo, Circus cf. C. cyaneus, Falco peregrinus/Falco biarmicus, Falco tinnunculus, Limosa sp., Larus sp., Columba livia, Tyto alba, Bubo insularis, Turdus sp., Sturnus sp., Corvus corax, Corvus corone/Corvus frugilegus, Pyrrhocorax pyrrhocorax, Pyrrhocorax graculus) (Malatesta and Suriano, 1970;Louchart, 2002;Louchart et al., 2005;Tyrberg, 2007). Malatesta (1970), discussed the stratigraphic setting and context of this fossiliferous deposit and hypothesized that it was contemporary to a fossiliferous layer outside the cave and that both likely dated to the last glacial event. ...
... The presence of the endemic eagle Aquila nipaloides, which is closely related to the present-day mainland species Aquila nipalensis that inhabits open dry environments (i.e. steppe-tundra, savannas and semi-deserts; Louchart et al., 2005), constitutes further evidence for savanna-like habitats. The Porto Conte area must also have been characterized either by ponds or small streams, as suggested by the occurrence of the remains of anserine and anatine birds, as well as of amphibians and reptiles, probably regurgitated in the vicinity of Medusa-Dragonara Cave by raptor birds (Kotsakis, 1980). ...
The timing of the colonization of Sardinia by mammalian fauna and anatomically modern humans (AMH) is currently under debate. The understanding of the geological and palaeoclimatological conditions that characterized the Late Pleistocene and the Holocene is essential to investigate colonization processes and requires an integrated multidisciplinary approach. In this research we combine for the first time new radiometric dating, field evidence of relative sea-level changes (RSL), sea-level and glacial-isostatic adjustment modelling, stratigraphical, palaeontological, palaeoecological and isotope analyses that were performed on sites and finds from the Porto Conte and Capo Caccia area (Alghero, NW Sardinia). The most important Late Pleistocene fossiliferous deposits in Sardinia have been discovered in this part of the island. Modelled palaeogeographical reconstructions of the Porto Conte area combined with the inferred environmental context and chronology of Late Pleistocene faunal assemblages from Medusa-Dragonara Cave suggest that Homo sapiens did not inhabit this area before the Holocene. Our results support the hypothesis that human presence in Sardinia during the Last Glacial Maximum was at best sporadic.
... The taxonomy was determined using the osteological collections of the Institut Català de Paleoecologia Humana i Evoluci o Social and the Museu de Ciències Naturals de Barcelona, as well as anatomical descriptions (Cohen & Serjeantson, 1996;Driesch & Boessneck, 1976;Louchart et al., 2005;Tomek & Bochenski, 2000). The remains were classified into three different weight classes considering the species identified in this study (Núñez-Lahuerta et al., 2021) (Table 1). ...
Bird assemblages can be found in archaeological sites throughout the Pleistocene. Taphonomy studies are key to understanding how such assemblages were formed. These assemblages can be generated by various agents, including human groups, animals and natural death. In this paper, we analysed the avifaunal assemblage from level TE9d at the Sima del Elefante site, where corvid (Corvidae) remains are the most abundant taxa, as reported in previous studies from that deposit. The remains of smaller birds (Charadriiformes, Rallidae) and large raptors (Aquila cf. heliaca/adalberti, Haliaeetus albicilla) have also been documented. Our taphonomic analysis shows that birds of prey were the main accumulators of bird remains at the site. The presence of modifications such as beak/talon marks and different degrees of digestion reinforce the idea that both diurnal and nocturnal raptors were involved. Nevertheless, we cannot rule out the accumulation by natural death of those species that could have nested in the rocky walls of the cave. The absence of evidence of anthropogenic activity on the bird remains from level TE9d therefore indicates that hominins were not involved in this accumulation. These results can be considered a new contribution to the discussion on the origin of avifaunal accumulations in Pleistocene archaeological contexts.
... grassy food or prickly bushes) and inhabiting similar environments (e.g. rocky hill), as suggested by De Vos and Van der Geer (2002)] had neither potential competitors nor predators, the impact of the endemic near-flightless (Athene cretensis) being negligible, and the presence of a second endemic raptor (Aquila according to Weesie, 1988, but Aves incertae sedis in Louchart et al., 2005) not substantiated because it is based on a single fragment, lacking diagnostic features. Accordingly, it is rational to suppose that inter-specific competition was low and individuals within each herbivorous species could efficiently exploit resources. ...
Islands have been regarded by scientists as a living laboratory of evolution, and a prime target for the study of forces influencing evolution. This research aims to investigate whether a dwarf deer (Candiacervus ropalophorus), that suffered a traumatic fracture of its metatarsal, might have survived, despite the broken limb, in the Cretan free-predator environment for a time longer than commonly expected on mainland. The metatarsal shows a healed complete and oblique fracture at level of the distal part of the diaphysis. Both macroscopic and X-ray examinations show the impressive new bone formation that occurred in the post-trauma period. The size of osseous callus, covering the fracture line and the surface next to the lesion, suggests that the deer has survived several months after the traumatic injury. In a mainland context, an injured deer would hardly have survived for such a long period, because any movement constantly irritated the fracture, dramatically reducing performance or willingness to move and the gait speed, and hampering any escape.
... Although avian carnivores tend to be more successful as island invaders than their mammalian counterparts, the only one certainly represented in the Quaternary of Crete is the endemic nearflightless owl (Athene cretensis). Whether there was a second endemic raptor is controversial: on the basis of a single fragmentary bone Weesie (1988) asserted that Aquila had once existed on the island, but as this element has now been referred to Aves incertae sedis (Louchart et al., 2005) there is no remaining basis for asserting the presence of an eagle. The Quaternary herpetofauna includes a number of small to medium-sized lizards, snakes, and turtles, including the extinct endemic tortoise Testudo marginata cretensis (Bachmayer et al., 1976). ...
Age-graded fossils of Pleistocene endemic Cretan deer (Candiacervus spp.) reveal unexpectedly high juvenile mortality similar to that reported for extant mainland ruminants, despite the fact that these deer lived in a predator-free environment and became extinct before any plausible date for human arrival. Age profiles show that deer surviving past the fawn stage were relatively long-lived for ruminants, indicating that high juvenile mortality was not an expression of their living a “fast” life. Although the effects on survivorship of such variables as fatal accidents, starvation, and disease are difficult to gauge in extinct taxa, the presence of extreme morphological variability within nominal species/ecomorphs of Candiacervus is consistent with
the view that high juvenile mortality can function as a key innovation permitting rapid adaptation in insular contexts.
Today, Sardinia and Corsica are two separate islands. Before the Holocene, however, they formed a unit with a shared geological history. Their fossil faunas are therefore practically identical. This chapter gives a concise overview of the faunas of Sardinia and Corsica. Then, a comprehensive treatment is provided of the history of discoveries, biozones, or faunal units and the peculiarities and evolutionary aspects of individual endemic species or lineages. With the start of the Miocene, the Sardo‐Corsican microplate separated from southern Europe and was inhabited by successive endemic insular faunas since then. During the Late Miocene, a temporary land connection with the island Tuscany resulted in a shared, insular endemic fauna. The most common elements are goral‐like bovids in the Miocene and Pliocene and dwarf megacerine deer and giant pikas in the Pleistocene.
This chapter gives a concise overview of the faunas of Crete. Then, a comprehensive treatment is provided of the history of discoveries, biozones, or faunal units and the peculiarities and evolutionary aspects of individual endemic species or lineages. The fossil fauna of Crete, the largest island of Greece, is mainly known for its Pleistocene endemic taxa, amongst which are dwarf deer, dwarf elephants and mammoths, dwarf hippopotamuses, and giant mice. The typical fauna elements of this biozone – the Mus zone – are two chronospecies of mice, Mus bateae and Mus minotaurus – unrelated to the Cretan mice of the previous biozone – Creutzburg’s dwarf elephant, Cretan deer, the Cretan otter, and the Cretan shrew. Excavations in the many cave localities – of which Michael Dermitzakis, Elliot Lax, and George Iliopoulos and colleagues gave overviews – revealed a clear zonation in the Pleistocene fossil record.
This Thesis is the result of the identification and study of part of the bird communities of the Upper Pleistocene and Holocene of the Pityusic Islands (Eivissa and Formentera) specially, Passeriformes and their potential predators (Falconiformes and Strigiformes).
All fossils studied come from a single site, es Pouàs (Sant Antoni de Portmany, Eivissa). The paleontological richness of this site is unique among the Mediterranean islands. The abundance of birds obtained in es Pouàs is related to the absence of terrestrial mammals in the Upper Pleistocene and Holocene of Eivissa.
Identifying bird fauna in es Pouàs was possible through the study and selection of thousands of fossils obtained in the excavation campaigns conducted between 1988 and 1994. Fossil identification was carried out using diagnostic features, both unpublished and obtained from comparison materials, as well as from scientific literature.
16,262 fossils have been identified belonging to 16 families of Passeriformes and 3 families of the orders Falconiformes and Strigiformes. Of a total of 73 taxa identified at a specific or generic level, 60 correspond to Passeriformes, 9 to Falconiformes and 4 to Strigiformes. Five species of raptors and 40 species of Passeriformes are included for the rst time in this site. There are 24 species of Passeriformes and six Falconiformes and Strigiformes which are new additions to the fossil record of the upper Pleistocene-Holocene of the Balearic Islands. We also have identified 28 species of Passeriformes not found in the continental fossil record near Eivissa.
In addition to studying the fossils of raptors and Passeriformes, the fossil fauna of es Pouàs was compared with the current ornithofauna of the Pityusic Islands as well as the current and fossil fauna of the adjacent area. The guild of birds of prey is the subject of a chapter in which both its bio-accumulator role as its uniqueness is valued.
The paleoecological and paleoclimatical significance of the bird fauna was studied.
The habitats that the identified species currently occupy allows us to better understand the environment of the Upper Pleistocene of the Pityusic Islands.
The presence of species with current northern distribution indicates the existence of a cooler climate in the Upper Pleistocene. Possible causes of local disappearances of some species and the absence of others from the fossil record are discussed. There are documented cases of morphological changes related to processes of insular evolution (e.g., Bubo sp., Corvus corax).
The study of the current diet of Tyto alba in the lesser Pityusic Islands was included. This study has allowed the evaluation of some of the effects of human arrival on these islands and provides an example of the usefulness of paleoecological studies in the interpretation of current communities.
Past human impacts on the riverine ecosystems of North America remain poorly understood. This continent is home to the world's most diverse freshwater mussel fauna, but mussels are particularly vulnerable to human impacts because they are long-lived sedentary filter feeders with complex life histories. A large body of historical and archaeological sources provide an extraordinarily comprehensive record of mussel distribution and in some cases abundance throughout the Holocene that exists for few organisms in general and is unprecedented for invertebrates. Despite high harvest pressure and the potential effects of prehistoric human land use practices on aquatic habitats, no extinctions of mussel species have been documented in North America until the 20th century. However, freshwater mussels have experienced one of the highest rates of extinction of any group of organisms during the past 100 years, primarily due to dam construction and the indirect effects of habitat fragmentation.
An extinct species, Athene angelis, larger than the other recent and extinct species of the genus Athene, is described from the Castiglione 3 and 1 localities, in Corsica. In the Middle Pleistocene of Corsica there is an association of three extinct Strigiformes, a dwarf Eagle Owl, Bubo insularis, a large Little Owl, Athene angelis, and a giant Barn Owl, Tyto balearica. There is an abridged English version.
The extinct owls from the Mascarene Islands can be ascribed to a distinct genus, Mascarenotus, probably derived from the genus Otus. Commerson's Scops Owl, from Mauritius, known from a description and a drawing, corresponds to the same taxon as "Strix' sauzieri, described from fossil bones. The bone remains recently found in La Reunion are ascribed to a new species, M. grucheti. The genus Mascarenotus shows a striking parallelism with the extinct genus Grallistrix, from the Hawaiian Islands. -English summary
Notes on the biogeography of the Corsican avifauna at the higher Pleistocène Holocene boundary, within a European context Changes in geographic range of bird species at the higher Pleistocene-Holocene boundary (10 000 years before present) are not all directly linked to global warming. In most cases regression or rarefaction were involved. The whole continent, as well as islands, was affected. Approximately 50 taxa, over 20 of which were part of the higher Pleistocene Corsican fauna, have been studied in relation to whithc factors caused their regression, disappearance or extinction at a Western Europe or Mediterranean islands scale. These factors were found to be mostly linked to climate changes, whereas during the Holocene human factors became dominant (destruction of species and habitats). The rapid regression followed by the disappearance of steppes or steppic tundra seems to be a recurring factor. Very cold events during the very end of the Pleistocene, a reduction in the emerged area of islands, the extinction of mammal species endemic to the islands, the intentional and unintentional introductions of mammal species by humans and human hunting were also identified as factors of some importance. All these factors interact and most of changes in distribution appear to be diachronic. Some groups such as Laridae, Sternidae and Paridae did not seem to be well established on islands before the Holocene, when a certain anthropophily started to develop. Islands, by their situation, were at first protected from some factors but were later found to be more venerable when exposed to these factors. Corsica and its rich Pleistocene avifauna were found follow this pattern and allowed us to define some modalities. Many species disappeared from Corsica following the retreat of steppic habitats and the simultaneous extinction of Megaloceros (Nesoleipoceros) cazioti this is clear in the case of carrion feeders such as the Magpie Pica pica and of the Vuture Gyps melitensis. Cold events followed by a drastic reduction in the area of emerged lands of the "Cyrno-Sardinian block" seem to have had an influence. Human impact, during this period, might have had an effect but this is hard to prove, a possible case of human influence in the Mediterranean being the extinction of Lesser White-fronted Goose Anser erythropus and another related species.
A preliminary study of the middle Pleistocene birds from Spina-gallo Cave (Siracusa, Sicily) shows an avifauna composed of 61 species (28 Passeriformes and 33 non-Passeriformes), including Anseriformes, Falconiformes, Gruiformes, Charadriformes, and Strigiformes. Three extinct taxa, probably new to science, include a large Tyto, a long-legged Athene, and a small species of Corvidae to be described later. Paleoenvironmental reconstruction of the site indicates a temperate climate, like the present or slightly colder.
1. In North America and México there is a strong tendency for island birds to have a longer tarsus and bill than their mainland counterparts; but there is no such tendency for them to have a longer wing and tail. 2. Although climate and body-size are known to influence the size of tarsus and bill the larger dimensions on islands are considered to be adaptations primarily to ecological conditions. The bill is longer because it deals with a greater range of food-sizes, and the tarsus is longer because a greater variety of perches is used. 3. The differences in usage have arisen as a result of the absence, on the islands, of species with similar ecological requirements, which has permitted some of those present to extend their activities and occupy at least part of the vacant niches, sometimes in new habitats. 4. It is suggested that the same ecological conditions and responses have influenced the evolution of large body-size of island rodents in particular, and of mainland animals of several phyla in general.