Fossil and subfossil birds of BrazilABSTRACT:
The study of fossil and subfossil birds in Brazil is still in its early stages despite its relatively abundant material. The remains are represented by bones, feathers, mummified specimens, eggs, coprolites, pellets, and tracks found in all Regions of the country starting in the 1830s. They are known from the Early Cretaceous to the latest Holocene, albeit several temporal gaps exist, and the most expressive diversity and quantity are concentrated in a few but important sites. Our survey and review of the literature and some previously unpublished specimens resulted in a list of 670 records demonstrating that the country’s known past avifaunas are essentially modern and rely mostly on provisional determination methods. Despite this, 15 extinct genera and 20 extinct species were newly described, demonstrating this field’s potential. The proper study of numerous long-stored materials from both paleontological and archeological contexts and eventual new remains can contribute significantly to our understanding of avian evolution and their taxonomic diversity throughout time, besides furnishing paleobiogeographic and paleoenvironmental information.
KEYWORDS: archeology; history of science; ornithology; paleontology; South America
INTRODUCTION
Brazil currently has 1,971 recognized recent bird species (Pacheco et al. 2021 ) and figures among the countries with the most expressive diversities in the world. While these recent taxa are particularly well studied (Sick 1997 ), knowledge of their prehistoric—fossil and subfossil (i.e., sub-recent, non-completely fossilized)—representatives has been progressing slowly, especially when compared with countries like Germany, China, the United States of America, and Argentina, the last holding the most complete and diverse known record in South America.
Taphonomic biases aside, several extrinsic factors contribute to this slow development of the study of avian remains in both paleontological (paleornithology) and archeological (archeornithology) contexts in Brazil (Alvarenga 1992 ; Queiroz 2010 ). The lack of reference osteological collections is a major obstacle, which perpetuates mentions of indeterminate avian material throughout the literature. The virtual absence of specialists dedicated to the study of fossil birds in the country is also a key factor, a scenario prompted by aspects such as a nationally generalized underappreciation of science and possibly the misleading idea that birds do not fossilize well and, therefore, not enough materials of this group exist for full-time dedication. There are several mentions of bird material in conference abstracts by undergraduate students, but, only in rare exceptions, this research line is carried on in their careers.
Nevertheless, the potential of this field in the country is demonstrated by the 20 extinct species that were formally described from paleontological localities across the country ( Fig. 1 ), with the oldest records dating back to the Early Cretaceous ( Fig. 2 ) (Winge 1887 , Ameghino 1891 , Alvarenga 1982 , 1983a , 1985a , 1985b , 1988 , 1990 , 1995, 1999a , Baird and Vickers-Rich 1997 , Olson and Alvarenga 2002 , Alvarenga and Guilherme 2003 , Alvarenga and Olson 2004 , Alvarenga et al. 2008 , Mayr et al. 2011a , 2011b, Carvalho et al.2015b, 2021 ). Overall, the remains are represented by bones, feathers, mummified specimens, eggs, coprolites, pellets, and tracks (e.g., Castro et al. 1988b , Alvarenga 1997 , Figuti 2005 , Marsola et al. 2014 , Lima et al. 2023 ) and are housed in museums or private collections in Brazil, Denmark, the USA, England, France, Germany, and Japan (e.g., Reinhardt 1881 , Winge 1887 , Ameghino 1891 , Olson 1981 , Alvarenga 1982 , Naish et al. 2007 ).
Map of South America with Brazil and its Federative Units showing noteworthy localities that yielded prehistoric material. Fm.: Formation; Ba.: Basin.
Simplified geochronological chart with the distribution of the most relevant extinct taxa with at least a generic attribution. Additional categories of material are also included. Numerical values in million years.
Published paleontological studies are focused almost entirely on taxonomy (Taranto 2012 ). Most of the material determined so far belongs to extant species, and yet its attribution to specific taxa cannot be accepted without a doubt (Olson 1985a). The literature is very decentralized, and older studies have rarely been considered in more recent ones. Several mentions are limited to lists of taxa without specifying the material’s nature and state. Most of the recent bird material described in major articles was analyzed by researchers who also study other groups (usually vertebrates) in their publications. Some studies also tend to be overlooked due to being published in volumes focused on different areas of prehistory knowledge.
Based on the above, this study aims to: (1) review the history of paleornithological studies in Brazil, focusing on the most important fossil-bearing localities; (2) catalog and integrate the data on paleontological and archeological records of Brazilian birds present in the literature; (3) update the nomenclature attributed to the materials following the most recent taxonomic and phylogenetic literature; (4) include descriptions and images of some unpublished material present in collections of Brazilian institutions; and (5) make available data originally published in non-scientifically universal languages, such as the pioneering Danish contributions.
The historical approaches published so far are incomplete, focusing mostly on the extinct taxa. The published material was listed, never in its entirety, in Lambrecht ( 1933 ), Brodkorb ( 1963 , 1964 , 1967 , 1971 , 1978 ), Mones ( 1986 ), and Cuello ( 1988 ). The last two are, in short, reproductions of secondary mentions, which, coupled with the fact that much of the published data is obscure, indirectly increases the possibility of misinformation dissemination.
The lack of an updated and comprehensive catalog of fossil and subfossil records is a significant impediment to our understanding of these taxa’s diversity and distribution over time, the type of material preserved, their paleoecology, and the overall state of knowledge of the national paleontological heritage. Remains from archeological sites are usually treated in zooarcheology focusing on the cultural context in which they were found. Since avian remains in Brazil are especially rich for the Pleistocene–Holocene transition and chronological control of these deposits is frequently incomplete, the exclusion of the archeological material would not be interesting from the perspective of taxonomic completeness.
The availability of information originally published in Danish (such as the studies of Peter Wilhelm Lund, Oluf Winge, and Johannes Reinhardt) also aims to solve a historical problem already mentioned by the Danish zoologist Theodor Mortensen ( 1925: 517) when he commented on Herluf Winge’s work: the lack of recognition and reach of ideas when not published in a universal language that, when they finally reach researchers from other parts of the world, are often either ignored or misinterpreted because of this language barrier, causing significant damage to science and the author’s recognition.
The gathering of data and the inclusion of unpublished information will provide a basis for further studies and in-depth reviews of the published material, offering new perspectives on the knowledge of these birds and their associated paleoenvironments.
MATERIAL AND METHODS
Both specialized technical literature and popular texts were consulted, along with previously unpublished documentary material. Undergraduate theses are not included in this review unless otherwise noted. Figures of the most important fossil taxa are presented, including some of historical significance that were only mentioned in the literature or depicted through images without good resolution. Credits of the figures are included in their respective subtitles.
Site names follow the original publications, with updated Portuguese spelling when necessary. All direct quotations are translated into English.
Some authors’ names that have been cited in different ways in the literature, especially regarding composite names (e.g., Cunha/Souza Cunha, Ribeiro/Mentz Ribeiro), hyphenated names (e.g., Martins-Neto/Martins Neto), and names with prepositions (e.g., De Queiroz/Queiroz), were standardized here seeking to maintain work and historical unit. In the Portuguese naming system, prepositions are generally excluded when a name is abbreviated or a surname is cited.
The codes for each of the Brazilian federative units used in this study’s catalog section are summarized in Table 1 . When a record has no clearly defined federative origin, the country code (BR) is used.
Data was gathered from June 2015 to June 2023 with occasional additional articles added up to June 2024.
The institutional abbreviations present in this study are summarized in Table 2 , including historical denominations. Non-recurrent abbreviations are explained in the body text.
The acronym BP stands for “before present”.
RESULTS
1. HISTORY
We can divide the work on Brazilian fossil birds into two main parts, the first conducted by Danish and the second by Brazilian researchers. However, this timeline is not continuous, as only a handful of studies were published in the first half of the twentieth century. In some scenarios (i.e., in Lagoa Santa and Taubaté and Bauru Basins), the paleornithological finds and descriptions are associated mostly with the action of individuals, while others were the result of decentralized combined efforts. The most important events are summarized in Table 3 .
The history of paleornithological research in Brazil was never explored in detail and was only partially summarized by a few authors over time.
In Denmark, Reinhardt ( 1881 ) commented on the fossil bird material gathered in the Lagoa Santa region by his fellow countryman, the naturalist Peter Wilhelm Lund, and related publications by other authors. While describing the collection gathered by Lund, Winge ( 1887 ) included a summary of what was published on the subject until then.
In Brazil, Goeldi ( 1894 ) reproduced and commented, in Portuguese, on the records of Winge’s study and updated some of his generic attributions based on the current literature. Padberg Drenkpol ( 1931 ) published and commented, in Portuguese, an extract of Winge’s study dealing with the paleontology of parrots. The compilation of Lund’s translated theses by Paula Couto ( 1950 ) included mentions of the bird material with comments. Valle and Carnevalli ( 1973 ) mentioned Lund’s research on fossil birds and his study on the relationship of barn owls with the formation of small bone heaps in caves. Ruschi ( 1979 ) summarized the finds, including mentions of zooarcheological material. Sick ( 1984a ) briefly mentioned the Paleogene finds from Rio de Janeiro and São Paulo, and Lund’s and Winge’s research (Sick 1984a , 1997 ), as well as subfossils and rock paintings depicting birds. He mentioned the occurrence of some fossil representatives in the texts in which he addressed the families. He further noted that the American paleornithologist Pierce Brodkorb provided most paleontological data, and, perhaps because of this, his coverage of Winge’s work is sparse, although he mentioned having a translated version of it. Alvarenga (1993a, 1997 ) briefly mentioned the earliest finds and focused on the material found in the country’s Southeast region, on which he was working. Alvarenga and Höfling ( 2000 , 2004 , 2011 ) highlighted most of the extinct taxa in their reviews of avian paleontology. Petri ( 2001 ), Ramos and Kellner ( 2005 ), Taranto et al. ( 2009b ), and Taranto ( 2012 ) briefly mentioned the extinct species, and Stefano et al. ( 2012 ) commented on the bird collections gathered by Lund, both living and fossil, although somewhat inconsistently, at times not specifying the nature of the mentioned material. Finally, the authors of the present work contributed with a thorough review of the history of the fossil material found by Lund and other researchers in the Lagoa Santa region (Nascimento and Silveira 2020 ) and an abstract of the present study (Nascimento and Silveira 2021 ). The present study is an expanded version of Rafael’s master dissertation (Nascimento 2022 ) and updates the authors’ previous works.
The country’s record of extinct fossil species was further mentioned, in various degrees, in general reviews that include South American fossil birds such as Tambussi and Noriega ( 1996 ), Mayr ( 2009 , 2016 , 2022 ), Tambussi and Degrange ( 2013 ), and Agnolín ( 2016b ).
Avian remains from the archeological record were never revised focusing on their taxonomical diversity.
The following sections explore the history of the most important fossil-bearing areas with bird remains in Brazil geochronologically. Information was gathered concerning the sites’ geological aspects and exploration history, emphasizing the discovery of bird material, paleoenvironmental aspects, and associated fossil taxa.
Summary of the institutional abbreviations used in this study. Note: this is not a list of the institutions housing Brazilian bird fossil material.
1.1. Crato Formation (CE)
1.1.1. General information
The Crato Formation ( Fig. 1 .1)—sensu Martill et al. ( 2007 ), Crato Member of the Santana Formation in some studies—outcrops around the northern, eastern, and southern-eastern flanks of the Chapada do Araripe plateau, at the boundaries of southern Ceará, western Pernambuco, and south-eastern Piauí. It comprises a heterolithic sequence of clastic and carbonate strata currently restricted within the Araripe Basin and some smaller, interconnected basins (Martill 2007 ). Martill and Heimhofer ( 2007 ) recognized four members in the Formation, beginning from the most basal: Nova Olinda, Caldas, Jamacaru, and Casa de Pedra. Only in the Nova Olinda Member, a series of laminated limestones up to 14 m thick, fossils occur in an astoundingly large number, often preserving exceptional details. The lowest part of the sequence of the Crato Formation, which includes the Nova Olinda Member, is presumably of late Aptian age in the Early Cretaceous (119–113 million years BP; Martill and Heimhofer 2007 , Carvalho et al. 2015b ).
The sediments of the Crato Formation were deposited in an equatorial mid-continental setting, with a hot and arid climate, in a lacustrine environment with a suggested minimum extent of about 18,000 km 2 (Martill 2007 , Carvalho et al. 2015b ). The water body was inferred to have considerably deep waters, with hypersaline events and anoxia in the bottom waters that assisted to some degree the exquisite preservation of the fossils by inhibiting efficient decomposition and disarticulation (Heimhofer and Martill 2007 ). However, it is noteworthy that these and other aspects remain debatable, and different approaches exist in the literature (see Martill and Bechly 2007 ).
Information on strata included in the Crato Formation was first published in 1846, but only by the 1940s and 1950s fossils began to be mentioned in the literature. It was not until the 1980s that serious paleontological research commenced, with numerous fossils found due to the significant increase in the extraction of limestones for commercial purposes (Martill and Heimhofer 2007 ).
Commercial mining for cement manufacture and limestones used as building stones, tabletops, and paving slabs occurs in the Nova Olinda Member and some of the overlying argillaceous strata, which makes it of considerable economic importance to the region (Andrade 2007 , Martill and Bechly 2007 ). Most fossils existing in scientific collections result from these commercial activities, including the subsequent illegal fossil trade. Quarry workers find the fossils while manually extracting the limestones and sell them at meager prices to middlemen based in Santana do Cariri and Nova Olinda in most cases, which in turn sell them to dealers based in São Paulo and abroad (Martill and Bechly 2007 ).
The Nova Olinda Member is an essential lagerstätte due to the diverse and abundant fauna and flora—with the most diverse non-marine fossil assemblage of Gondwana (Martill et al. 2007 )—and for the exceptional quality of preservation, with a high level of anatomic fidelity, mostly in three dimensions, and many unweathered specimens retaining color patterns (Martill and Frey 1995 ). The fossil assemblage is dominated by arthropods, with insects being the most abundant group in both number and taxonomic diversity, but also represented by chilopods, arachnids, and crustaceans. The vertebrates include fishes, with the gonorynchiform Dastilbe crandalli being the most abundant, anurans, turtles, lizards, crocodilians, and pterosaurs (Martill et al. 2007 ). Besides birds and indeterminate feathers, the only other known dinosaur record is the compsognathid not properly described but known by the unavailable name of Ubirajara jubatus (Smyth et al. 2020 ), which preserved filamentous integumentary structures. The small, volant birds known from the locality may have been responsible for predation marks observed in some fossil insects (Naish et al. 2007 ). The fossil flora is diverse, dominated by gymnosperms (such as conifers, cycadophytes, and gnetophytes) and angiosperms, but also includes several spore-bearing plants (Mohr et al. 2007 ).
1.1.2. Feathers
In the Crato Formation, remains of possible birds are extremely rare and were so far reported only from the Nova Olinda Member (Naish et al. 2007 ). Small, isolated feathers were likely blown into the Crato lagoon, where they sunk and were rapidly buried in the anoxic bottom (Martill and Davis 2001 ). They occur predominantly as carbonized traces, though melanosome and possibly iron oxide preservation are also present (Prado et al. 2016a ). Apart from being categorized by comparison with modern feather morphotypes, these specimens could not be assigned to any taxon. They possibly represent several different and disparate taxa, and not even their origin from birds can be secured since some non-avian theropod groups also possessed true feathers (Naish et al. 2007 ). However, after their association in one case with avian skeletal remains (Carvalho et al. 2015a , 2015b ), Smyth et al. ( 2020 ) postulated that most are likely from enantiornithiform ornithothoraceans rather than from non-avian dinosaurs. The deposit represents the only case of fossil feathers associated with skeletal remains where feathers occur frequently in South America. It is also one of the few localities that bear Mesozoic bird remains in Gondwana. Apart from the Yixian Formation in China, where complete avian fossils associated with feathers are common, it is one of the richest Mesozoic feather localities (Naish et al. 2007 ) and constitutes the only known record of feathers of this age from Brazil (Metello 2017 ).
The first-ever fossil described as Mesozoic avian remains from Brazil was one of these isolated feathers, a probable remex associated with a well-flighted bird by Martins Neto and Kellner ( 1988 ). This record was also the first description of a fossil feather from South America. Its exact origin is unknown, but it was probably collected around Santana do Cariri, Ceará. It is preserved as a limonitic by-product of a weathered pyritic permineralization (Martill and Filgueira 1994 ). This feather’s discovery in a site that provided small, well-preserved vertebrate fossils (e.g., fishes and anurans) along with other fossils (such as insects and seeds) indicating abundant and varied food sources offered prospects for future discoveries of more complete bird fossils (Kellner et al. 1991 ). Referring to this publication, Alvarenga and Höfling ( 2000 , 2004 ) commented that the feathers from this deposit most likely belong to Enantiornithes, without presenting further information to support this suggestion.
The second feather, this time a semiplume, was reported by Martill and Filgueira ( 1994 ), being overall the third reported occurrence of Mesozoic avian remains from Brazil. The fossil comes from the then recently discovered locality of Mina de Antone Phillipe, near the Tatajuba Reservoir and about 10 km west of Nova Olinda, Ceará, Crato Formation’s main commercial exploitation area. One of the quarry workers collected it and passed it down to Antone Phillipe, the quarry owner, as a fortunate event despite being on a site outside the main commercial collecting area, where most fossils were ignored. The fossil appears to have been preserved organically, although no tests were conducted for fear of damaging it further. The Crato Formation at this locality was probably deposited close to the ancient shoreline of the Crato lagoon.
Kellner et al. ( 1994 ) described a fossil down feather without attributing it to a specific locality. Nevertheless, the record was considered empirical evidence that the Early Cretaceous birds had already developed an effective thermoregulatory insulation cover and increased the expectation of finding skeletal remains of birds in the same deposit.
Martill ( 1994 ) pointed out fake fossil feathers among the fabrications done by local quarry workers intended to supply the illegal fossil trade. Modern feathers were glued to limestone slabs, with some fakes challenging to detect. He purchased two specimens ( Fig. 8 in the article) that could not be determined as fabrications in the field without a good microscope. Their slight elevation on the bedding plane and the fact that one slab had two feathers in suspiciously similar orientations were clues to their possible fake identity.
Martill and Frey ( 1995 ) reported feathers from exposures of unweathered, gray-colored laminites at the Mina de Antone Phillipe. One of them, an incomplete contour feather with alternating dark and light bands, was regarded as perhaps the earliest recorded occurrence of color patterning in birds. The authors deposited part and counterpart of the specimen in different institutions aiming for a wider availability for examination. Several other non-flight feathers were collected on the same field trip in which this banded feather was obtained. These feathers are preserved as organic material of a dark appearance. Martill and Frey did not risk examining the feather’s color pattern (as in insect specimens with the same kind of preservation) by electron microscopy as the pattern preservation could be masked by gold or carbon coating. Instead, they chose to analyze another feather from the same locality with a similar style of preservation despite lacking the color patterning. At high magnifications, it showed that the feather’s organic material is composed of elongated and roughly aligned rod-shaped bodies that the authors considered autolithified bacteria similar to material described from the Eocene oil shales of Messel, Germany.
Martill and Davis ( 1998 , 2001 ) described an almost symmetrical feather with possible ectoparasite eggs. The specimen was obtained in June 1996 from a commercial fossil dealer by the National Science Museum of Japan, and, although the locality was given only as Crato Formation, its matrix is rather similar to that of weathered, laminated limestones of the Nova Olinda Member, and was probably collected in one of the small quarries operated between Nova Olinda and Santana do Cariri. It is too weathered for its original preservation mode to be satisfactorily established. The numerous spherical, hollow structures about 75 μm wide were interpreted as probable feather mites and regarded as possibly the oldest occurrence of ectoparasitism on feathered maniraptorans. It is the longest feather known from the Crato Formation and a little larger than all Cretaceous feathers reported until then, with a length of 85 mm.
Kellner and Campos ( 2000 ) noted that several feather morphotypes had been unearthed in the Formation, most of which remain undescribed. They mentioned about two dozen of unknown provenance. Although they suspected all published specimens might have come from outcrops around Nova Olinda (as several specimens were, in any case) since in the past all the mining sites in the Formation were in that area, the expansion of the quarrying activities in the region led to the opening of several new mining pits around Santana do Cariri and along the Nova Olinda–Santana do Cariri road. Thus, some of the newer specimens might have come from them. Regarding their identity as avian or non-avian dinosaurs, they considered that down and flight feathers likely belonged to birds. “True downs” were not known in non-avian dinosaurs, and a strongly asymmetrical remex was more consistent with a well-flighted animal—though Bittencourt and Langer ( 2011 ) noted that the occurrence of asymmetrical feathers in non-avian eumaniraptorans challenges this attribution. The nature of the other morphotypes remained inconclusive, but they also observed that they did not know about semiplumes occurring in non-avian forms. The development of distinct coloration patterns known on some contour feather specimens was also highlighted, which shows that artifices observed in modern species (such as behavioral and communicational) were likely present or had similar variants in this stage of dinosaur evolutionary history.
Kellner ( 2002 ) described a complete contour feather preserving a banded color pattern, previously depicted by Kellner and Campos. Additionally, he mentioned that several very small feathers (less than 20 mm), either down feathers or semiplumes, are housed in the American Museum of Natural History, New York, and MN.
Semiplumes frequently occur in the Crato Formation, with Naish et al. ( 2007 ) having examined some 20 or more, compared with only one possible remex and two or three symmetrical rectrices. These feathers are mostly small, 10–20 mm long, but some larger ones are also present, with one reaching 22 mm.
Vinther et al. ( 2008 ) analyzed the part specimen of the banded feather originally described by Martill and Frey ( 1995 ) in a study about the color of fossil feathers, along with Eocene material from the Fur Formation of Denmark. By comparing the structure of black feathers of a living icterid, they interpreted the elongate, oblate carbonaceous bodies making up the dark bands of the Crato feather as eumelanin-containing melanosomes. Their observations indicated that structures present on fossil feathers previously reported as bacteria (as the second feather mentioned by Martill and Frey) are melanosomes, of which the distribution can preserve the color pattern of the original feather in the fossil. Thus, this study proposed that exceptionally preserved melanosomes were a more plausible scenario as the chief explanation for integumentary preservation in vertebrate fossils rather than lithified bacteria (Smithwick and Vinther 2020 ).
A down feather and a semiplume collected at the Mina do Demar, on the road connecting Nova Olinda and Santana do Cariri, during the field season of 2009 by biology student Laiz Karla of CAV were briefly described by Sayão and Uejima ( 2009 , 2010 ) and later depicted by Sayão et al. ( 2011 ). Additionally, Sayão et al. ( 2011 ) described a third feather with a morphotype like that of down or ornamental feathers from the same locality. They also speculated on the nature of the covert feathers in the deposit, relating their number, besides taphonomic aspects, to molting processes, and framed the described morphotypes to the developmental stages II (the third feather from Mina do Demar), III (semiplumes and down feathers) and possibly V (primary remex) defined by Prum ( 1999 ), still not confirming the presence of Aves in the deposit. França et al. ( 2017 ) provided measurements for the feathers described by Sayão and Uejima and other five specimens.
Leite and Hessel ( 2011 ) reported eight seemingly contour feathers (an isolated specimen and a set of seven feathers in the same block) from Mina do Triunfo in Nova Olinda. The set is unique, as it is the only published example of this type of conservation without associated osteological remains. Based on similarities with the fossil feathers from the Early Cretaceous of Liaoning, China described by Zhang et al. ( 2006 ), the authors tentatively attributed the specimens to small non-avian dinosaurs, possibly contour feathers from the upper part of the hindlimbs. Campos et al. ( 2019 ) analyzed the set of seven feathers through combined microscopy and spectroscopy techniques, aiming for minimal damage to the specimen. Their result confirmed the feather identity of the fossil and revealed eumelanosomes in its composition that indicate the feathers were originally dark-colored.
Prado and Anelli ( 2013 ) briefly described a semiplume and a down feather. These fossils were apprehended by the Polícia Federal and the Instituto do Patrimônio Histórico e Artístico Nacional (IPHAN) and deposited in the collection of the Laboratório de Paleontologia Sistemática of IGc/USP. The authors noted that the different feather morphotypes could represent ontogenetic stages or be interpreted as signs of intraspecific and behavioral relationships, such as the presence of neonates or parental care. Later, Prado et al. ( 2016a ) described these specimens further, classifying both as down feathers and assigning them to Coelurosauria, and presented an additional semiplume with the same apprehension origin, referring to it as Maniraptoriformes. They also developed on the taphonomy and paleoecology of the animals that provided the fossil feathers, speculating that the specimens initially described by Prado and Anelli were possible auricular feathers of a chicken-sized animal, while the third feather had both protective and thermoregulatory roles. Furthermore, possible roles in camouflage, communication, and sexual selection were suggested. They also speculated that birds might have lived by the shoreline of the Crato lagoon, which enhanced the probability of preservation of their remains. Finally, they developed on the absence of feathered dinosaurian remains in the Romualdo Formation [1] (the second, younger lagerstätte in the Araripe Basin known for preserving soft tissues such as muscle fibers and blood vessels), which was possibly related to different taphonomic conditions. Overall, the IGc/USP collection has 67 Crato feathers (some preserved as part and counterpart) with origin as apprehensions (G.M.E.M. Prado, personal communication).
Additionally, Prado and Anelli ( 2015 ) performed SEM analysis on two feathers, one from the Crato Formation (GP/2E-8771) and the other from the Tremembé Formation (GP/2E-8125). They found oblate microbodies in both fossils (not occurring in the matrix), interpreted as fossilized eumelanosomes. Most are degraded but indicate a high density and organization, varying in size from 500 to 1800 nm, morphologically suggesting an association with an iridescent pattern. By the known environmental settings from both sites, dry and arid for the Crato and hot and humid for the Tremembé Formation, they suggested dinosaurs from both units had dark or iridescent feathers, being well adapted to the ecological niches they occupied. Prado ( 2017 ) developed on these feathers further and analyzed them geochemically and microscopically along with specimens from the Tremembé Formation. He described their taphonomy (physical and molecular), ultrastructural elements (such as melanosomes and minerals), and preservation process.
Metello ( 2017 ) analyzed the plumage of Cratoavis cearensis (see below) and described all the 45 isolated fossil feathers in the UFRJ collection (though he depicted only six), which constitute 28 contour feathers, 8 plumes, and 9 semiplumes [2] , preserved as carbonized traces in all cases except for one preserved as an impression with a faded carbon contour. These specimens were collected between 2000 and 2016 in Mina Pedra Branca in Nova Olinda, Ceará.
Fourteen fossil feathers were mentioned by Nascimento and Oliveira ( 2018 ) and fully described and depicted by Nascimento ( 2020 ), consisting of two from the UFRPE collection, five from MPSC of URCA in Santana do Cariri and seven from the Laboratório de Paleontologia of URCA in Crato. They were prepared and determined as five plumes, three semiplumes, and six contour feathers, all showing good preservation with identifiable rachises, barbs, and barbules, two with banded color patterns.
Other isolated reports of fossil feathers include that of Teixeira and Saraiva ( 2010 , 2011 ), Lopes et al. ( 2017 ), Bantim et al. ( 2022 ), and Sousa et al. ( 2023 ). Several other unpublished examples are held in national and international collections, public or private (G.M.E.M. Prado, personal communication). Here we briefly present 67 feathers from the IGc/USP, LEG, and MHNC collections (see below).
1.1.3. Skeletal remains
Skeletal remains were first mentioned and depicted in the literature by Naish et al. ( 2007 ), although these authors stated that this kind of fossil was already known from anecdotal accounts and personal observations. Two specimens were preliminarily described, the first consisting of feathers associated with presumed carpal bones and deposited in the Senckenberg Museum in Frankfurt, Germany. The second consists of a poorly preserved partial skeleton with associated feathers, held in the Masayuki Murata (MURJ) private collection in Kyoto, Japan. By 2006, Martill and four other researchers were disputing the private museum’s permission to describe the fossil, considered the oldest Gondwanan avian fossil known from skeletal remains (if indeed representing a basal bird), but only photographs became available. It reportedly reached Japan through Germany and was sold for US$ 100.000 (Amorim 2006 ).
Herzog et al. ( 2008 ) reported rumors of a nearly complete and excellently well-preserved avian skeleton sold and smuggled out of the country, with no information about its destination. It is unclear whether this refers to the MURJ specimen (as they do not cite Naish et al.) or another trafficked specimen. The first seems more likely.
A fully articulated skeleton associated with feathers of a very small euenantiornithean bird was first described by Carvalho et al. ( 2015a ) and further by Carvalho et al. ( 2015b ), who classified it as the new genus and species Cratoavis cearensis . It is the first Mesozoic bird named from Brazil and the Early Cretaceous of South America and the most complete avian specimen of this age known from Gondwana. The fossil was found in Mina Pedra Branca by Cleuduardo Laurentino Dias, Devânio Ferreira Lima, and Antonio Josieudo Pereira Lima, who provided it to the authors to be studied. The specimen preserved long, rachis-dominated rectrices in relief, which shed light on the anatomical structure and probable function of this feather morphotype, previously known only from two-dimensional slabs and unknown among living birds, assisting, for example, in the determination of fossil material with complicated history as a true feather of this kind (Agnolín et al. 2017b ). Some color patterning was also possibly preserved in a row of lunar transversal bands in the tail feathers that, despite the specimen’s possible juvenile nature, resemble the feathering of modern adult birds. A South American record of this type of tail feathering also broadened this morphotype’s paleobiogeographic distribution, which was only reported until then from China.
In 2020, it was reported that researchers at the LACEV (Laboratorio de Anatomía Comparada y Evolución de los Vertebrados) of MACN were working on new contributions on the Early Cretaceous birds from Brazil (SAPE 2020 ). In the next year, Carvalho et al. ( 2021 ) described as an ornithuromorph the new genus and species Kaririavis mater from an isolated foot found in Mina Pedra Branca, adding to the otherwise enantiornithean-restricted avifauna of the Crato Formation. This small bird possibly belongs to an unknown clade with some cursory similarities with extant flightless ratites and represents the oldest known ornithuromorph from Gondwana. This record in South America demonstrates that basal ornithuromorphs were present and probably diversified in the Southern Hemisphere by the Aptian.
1.2. Bauru Group (MG, SP)
1.2.1. General information
The Bauru Group (sensu Soares et al. 1980 ) is the most widespread continental rock unit with vertebrate remains of Cretaceous age (Turonian–late Maastrichtian) in South America, being exposed over an area of 240,000 km 2 in western São Paulo, western Minas Gerais, northwestern Paraná, eastern Mato Grosso do Sul, and southern Goiás (Bertini et al. 1993 , Candeiro 2005 , Bittencourt and Langer 2011 , Candeiro et al. 2013 ). Different views on its geological evolution and stratigraphy exist in the literature, and, especially due to difficulties to refer earlier fossil finds to new units of more elaborated stratigraphic frameworks, most paleontological works still rely on a more traditional nomenclature (Bittencourt and Langer 2011 , Langer et al. 2022 ). Accordingly, here we aim to contextualize the terminology associated with avian fossil finds.
Four formations proposed by Soares et al. ( 1980 ) are traditionally recognized as constituting the Bauru Group as part of the Paraná Basin (in ascending order): Caiuá, Santo Anastácio, Adamantina, and Marília, with the last two richly fossiliferous (Bertini et al. 1993 ). The view of these deposits as an individualized Bauru Basin was proposed by Fernandes and Coimbra ( 1996 ), who erected the Caiuá Group, divided into Santo Anastácio, Rio Paraná, and Goio Erê Formations, and the Bauru Group, divided into Adamantina, Uberaba, and Marília Formations. A stratigraphic revision by Fernandes and Coimbra ( 2000 ) divided the Bauru Group into the Uberaba, Marília, Vale do Rio do Peixe, Araçatuba, São José do Rio Preto, and Presidente Prudente Formations (the last four once making up the former Adamantina Formation). Avian material has been associated in the literature with the Adamantina (and its dismembered formations) and Marília Formations of São Paulo and Minas Gerais.
The Adamantina Formation ( Fig. 1 .2) is generally regarded as Campanian–Maastrichtian in age (Gobbo-Rodrigues et al. 1999 , Santucci and Bertini 2001 ), but other timespans, such an older, Turonian–Santonian (Dias-Brito et al. 2001 ) or Santonian–Campanian (Langer et al. 2022 ) age, have also been suggested. Nevertheless, it originated in different depositional cycles and some faunal elements suggest it congregates rocks of significantly different ages with still poorly sampled individual biotas (Bittencourt and Langer 2011 ). The unit is predominantly composed of reddish clays and sands of eolian, lacustrine, and fluvial origin, formed under a warm, seasonal, and arid to semi-arid climate intercalated with more humid intervals (Bertini et al. 1993 , Bittencourt and Langer 2011 ). It is exposed in Goiás, São Paulo, and the Triângulo Mineiro region of Minas Gerais (Candeiro 2005 , Candeiro et al. 2013 ).
The Marília Formation ( Fig. 1 .3) is generally attributed to the Maastrichtian (Dias-Brito et al. 2001 , Candeiro 2005 , Bittencourt and Langer 2011 ) and is divided into three members: Ponte Alta, Serra da Galga, and Echaporã, the first two restricted to the Triângulo Mineiro region and the last exposed in Goiás and São Paulo (Candeiro et al. 2013 ). It is dominated by whitish calcareous conglomeratic sandstones of fluvial origin, deposited under relatively drier conditions (Bertini et al. 1993 ). Despite occurring over a large geographic area, the Adamantina and Marília Formations present very few natural exposures, with site-specific and restricted fossil localities. Most fossils were collected during the construction of roads and railways and excavations of buildings and wells (Bertini et al. 1993 ).
Vertebrate fossils from the Bauru Group were first reported in 1911, with small-sized vertebrates only extensively recovered at the end of the 1980s (Bertini et al. 1993 ). It contains one of the richest Late Cretaceous vertebrate assemblages of Brazil, including theropod and sauropod dinosaurs, crocodilians, turtles, and fishes, with relatively rarer occurrences of anurans, squamates, and mammals (Candeiro 2005 ). Invertebrate remains include bivalves, gastropods, crustaceans, and worm burrows and tubes, and the fossil flora includes charophytes, spores, gymnosperms, and angiosperms (Bertini et al. 1993 ).
Bird remains are uncommon in the Bauru Group (Azevedo et al. 2007 ). Chiappe ( 1991 ) mentioned an unpublished fragmentary proximal end of an avian carpometacarpus from the Bauru Formation through communication with Herculano Alvarenga in 1988 . Later, Chiappe ( 1996 ) was informed by Alvarenga that the Cretaceous material was contaminated, and the bone belonged to the extant Coccyzus americanus . Truly fossilized avian material was only discovered beginning of the next decade in localities of São Paulo and Minas Gerais.
1.2.2. São Paulo
The first true Mesozoic bird osteological remains from Brazil were presented to the public on 10 August 2005 during the II Congresso Latino-Americano de Paleontologia de Vertebrados in Rio de Janeiro. They were found just two months before in a site of 2 x 1 m and 50 cm deep discovered in September 2004 by the paleontologist William Roberto Nava (MPM) ( Fig. 3 A), who studied them along with Herculano Alvarenga (MHNT) (Castilhos 2005 , Thomé 2005 , Chiappe et al. 2018a ). Alvarenga and Nava ( 2005 ) reported the occurrence of three to four different sparrow-sized enantiornithean taxa from the Adamantina Formation of Presidente Prudente municipality, based on dozens of well-preserved, isolated, and partially articulated bones showing no evidence of reworking. The material is very similar to Enantiornis leali from the Late Cretaceous of El Brete, Argentina, but no new taxa were named due to difficulties in associating the bones. The uncertain locality of the site, however, does not allow attribution to the Vale do Rio do Peixe or the Presidente Prudente Formations of Fernandes and Coimbra ( 2000 ) (Langer et al. 2022).
Adamantina Formation, the work of William Roberto Nava: Nava in 2010 during one of the excavation phases of new small enantiornithean bones (A); plaque in the fenced area of William’s Quarry in 2022, two years after protection was granted by the municipality of Presidente Prudente (B) (W.R. Nava).
More material was discovered in hundreds of excavation trips—nearly 1,000 bones of dozens or even hundreds of individuals of small to medium-sized taxa (Chiappe et al. 2018a , 2018b , 2019 , 2022 , Fonseca 2019a ). In 2015, during a paleontological event in Uruguay, Nava and Argentinean paleontologist Luis M. Chiappe (LACM) formed a partnership to study the material, with Alvarenga and Argentinean paleontologist Agustín G. Martinelli (MACN) also contributing. Chiappe visited the region two times in 2017 and a third time in 2018 along with American fossil preparator Maureen Walsh and illustrator and photographer Stephanie Abramowicz (Prefeitura de Marília 2018 ). He returned in 2019 when more material was found in the fourth excavation phase (the second with Argentinean and American collaboration), which also integrated technician Guillermo Aguirrezabala and doctorate student Sebastián Rozadilla (MACN), as well as technician Jonatan Kaluza (Fundación Félix de Azara) from Argentina (Fonseca 2019a , Tomazela 2019 ).
The site, located in an urban public plot in the district of Parque dos Girassóis, was first dubbed Enantiornithes outcrop (Nava 2012 , 2013 , 2015 ) and later William’s Quarry (Chiappe et al. 2018a , 2018b , 2019 ), with the fossils concentrated in a very small area of about 6 m 2 of red-pink fluvial sandstones and claystones, possibly as the result of a catastrophic event (Nava in Lopes 2020 ). It also provided, in smaller numbers, remains of other small vertebrates such as fishes, anurans, chelonians, squamates (including the new lizard Brasiliguana prudentis ), and teeth of mesoeucrocodilians and theropod and sauropod dinosaurs, besides gastropods and coprolites (Nava 2012 , 2013 , 2015 , Nava et al. 2015 , Fonseca 2019a ). The site constitutes the most abundant Mesozoic avian locality in the Americas and the richest of Late Cretaceous age globally. It provides critical information for contrasting hypothesis of avian diversification during the Cretaceous–Paleogene transition and the earliest divergences of modern birds, and indicates, along with other contemporary Gondwanan localities, a notable abundance of enantiornitheans during the interval of 80–70 million years BP, which is difficult to reconcile with the hypotheses of a Southern Hemisphere origin for neornitheans during this time (Chiappe et al. 2018a , 2018b , 2019 ).
The site also provided the first confirmed examples of toothed birds from the Late Cretaceous of South America, adding important information to its otherwise meager cranial anatomy knowledge (Nava et al. 2015 ). Wu et al. ( 2019 , 2021 ) μCT scanned at high resolution two premaxillae and a partial dentary aiming for a better understanding of the occurrence of polyphyodonty in Mesozoic birds. The resulting data show an alternating dental replacement pattern in the premaxillae that is consistent with the observed pattern among extinct and extant reptiles, and dental replacement at different stages is revealed by the dentary. These data strongly suggest that an alternating pattern was typical of enantiornitheans and imply that the tooth cycling control mechanism is conserved during the transition between non-avian reptiles and birds.
Chiappe et al. ( 2022 ) performed the first detailed study of an enantiornithean endocranium on a nearly undistorted fossil found in William’s Quarry that presented traits traditionally regarded as exclusive to Neornithes, such as an expanded, flexed brain, a ventral connection between the brain and spinal column, and a modified vestibular system. This discovery suggests these endocranial characters may have originated before the split between Enantiornithes and the more crownward portion of avian phylogeny over 140 million years BP.
Paixão et al. ( 2023 ) developed on the taphonomic aspects of William’s Quarry focusing on the enantiornithean remains. They divided their study area into two points, PWQ01 (bonebed 1) and PWQ02 (bonebed 2), encompassing different levels and taphonomic processes. Material from PWQ02 presented a higher degree of articulation and orientation than that of PWQ01. Avian remains were more concentrated in PWQ01. Apart from those possibly belonging to Crocodilyformes and Theropoda, small and thin eggshell fragments that could belong to Enantiornithes were found at the site, but they are still under study (G.M.X. Paixão, personal communication).
Due to its importance, Nava and the associated researchers first requested the Presidente Prudente city managers to protect the site during the first phase of collaborative excavations in May 2017 (Fonseca 2019a , 2019b ). The protection was granted in a municipal decree on 14 February 2020 and the site area was fenced on 28 July 2020 ( Fig. 3 B), with the city intending to transform it into a paleontological park (Prefeitura de Presidente Prudente 2020 ).
The material from William’s Quarry was not the only one found in the State. Azevedo et al. ( 2007 ) reported a phalanx fragment of a possible bird among microvertebrate material collected through screenwashing technique at the Jales locality in the Fazenda Furnas of Adamantina [Vale do Rio do Peixe] Formation in Alfredo Marcondes (Candeiro 2015 , Alves et al. 2016 ), which shows sandstone sediments that are reddish, sometimes conglomeratic, and cemented by calcium carbonate. The fossil was collected in 2001 during an expedition led by paleontologist Lílian Paglarelli Bergqvist (UFRJ).
Marsola et al. ( 2012 ) reported the first Mesozoic avian egg from Brazil, found in July 2011 in a well-defined intraformational conglomerate level exposed within white-reddish sandstones attributed to the Adamantina [Vale do Rio do Peixe [3] ] Formation along road SP-270, near Álvares Machado. The nearly complete specimen was attributed to a basal Ornithothoraces and is one of the smallest known Mesozoic bird eggs (Marsola et al. 2012 , 2014 , Marsola 2013 ).
Other studies include Cardia ( 2019 ), which featured bird remains in the Bauru Group vertebrate material analyzed for mercury concentrations aiming at trophic chain reconstructions. Taranto et al. ( 2010 ) mentioned bird remains among the material collected from 2002 onwards in the Presidente Prudente Formation of Flórida Paulista and Alfredo Marcondes. Pinheiro et al. ( 2019 ) mentioned bird material among fossils collected in Presidente Prudente Formation (likely in William’s Quarry) during fieldwork organized by the Faculdade de Formação de Professores of UERJ in 2019. Lopes et al. ( 2013 ) reported a small (3 x 5.5 mm), conical tooth (UFRJ-DG 500 R (d)) from the Marília Formation of Alfredo Marcondes that shows features commonly observed in enantiornitheans and represents a new morphotype for the Bauru Group, but its identification was not conclusive.
1.2.3. Minas Gerais
The first record of Cretaceous bird material from Minas Gerais was that of Candeiro et al. ( 2010 ), who reported an isolated pedal phalanx of an indeterminate Aves of late Maastrichtian age. It comes from the Serra da Galga Member of the Marília Formation of Uberaba in the Triângulo Mineiro region, which contains one of the richest Late Cretaceous continental biotas in Brazil (Candeiro et al. 2013 ). Candeiro et al. ( 2012 ) reported an ungual phalanx of indeterminate Aves and a metatarsal of a possible Enantiornithes. These specimens were found during screenwashing performed by the CPPLIP starting in 2009 in the Ponto 1 do Price site of Peirópolis locality. The material is similar in degree of incompleteness to the material from the Adamantina Formation of Jales locality in São Paulo. The Peirópolis area provided a diverse Late Cretaceous fauna, including bivalves, ostracods, fishes, turtles, lizards, mesoeucrocodylians, and theropod and sauropod dinosaurs. Rédua et al. ( 2016 ) hypothesized these birds possibly fed on small freshwater or terrestrial invertebrates.
On the lack of avian records from the Adamantina Formation in the Triângulo Mineiro region, Candeiro ( 2015 ) pointed out this is probably associated with the lack of more intensive prospects in that unit.
1.3. São José de Itaboraí Basin (RJ)
1.3.1. General information
The São José de Itaboraí Basin ( Fig. 1 .4), discovered in 1928 by engineer Carlos Euler, is in the São José neighborhood of the district of Cabuçu in Itaboraí, Rio de Janeiro, about 60 km from the State’s capital. It is one of the smallest Brazilian sedimentary basins, with an approximate length of 1.4 km (northeast-southwest) and a minor axis width of approximately 0.5 km (northwest-southeast), with an area of approximately 1 km 2 . It is, however, richly fossiliferous, being filled by a sequence of clastic and chemical (travertine) limestones that are cut vertically by dissolution channels filled with marl, in which most fossils were found—avian remains are known from both gray limestone and channels (Bergqvist and Almeida 2005 ). The basin’s limestone was exploited between 1933 and 1984 for the manufacture of cement used in constructions such as the Maracanã Stadium and the Rio–Niterói Bridge, and this exploitation was responsible for the discovery of many fossils. The area, originally owned by the Companhia Nacional de Cimento Portland Mauá, was expropriated in 1990 by the municipality of Itaboraí, which created in 1995 the “Parque Paleontológico de São José de Itaboraí”, aiming at its preservation and the dissemination of its geo-paleontological importance (Bergqvist et al. 2006 , 2009 ). Although classically considered to be of late paleocenic age, the age attributed to the basin is now accepted as the early Eocene (Itaboraian SALMA, 53–50 million years BP; Gelfo et al. 2009 , Woodburne et al. 2014a , 2014b ). In addition, pleistocenic fossils and archeological materials have been found in the basin’s vicinity (Bergqvist et al. 2006 , 2009 ).
1.3.2. Avian remains
The first publication on the basin’s fossils appeared as early as 1929 and addressed incomplete internal molds of gastropods (Bergqvist et al. 2006 ). In 1948, the naturalist Ney Vidal, from the former Divisão de Geologia of MN, collected some fragments of bird bones there. Later, in 1949, Júlio da Silva Carvalho collected in the same place most of the components of a small collection of sparse and fragmented bird bones, which were included in the vertebrate collection assembled by the Seção de Paleontologia of DNPM (Alvarenga 1983a ). These fossils were briefly mentioned by Paula Couto ( 1953 , 1958a , 1970 ), who mentioned several unidentified birds, Palma ( 1973 ) and Palma and Brito ( 1974 ), who mentioned long bone fragments, Price (in lit. 1977 in Sick 1984a ), who mentioned the remains of several birds (“metatarsals, vertebrae, premaxillae, etc.”), Francisco and Souza Cunha ( 1978 ), who also mentioned long bones, and Rich ( 1979 ), who mentioned small birds. However, no detailed description would be published until Déa Regina Bouret Campos, then responsible for the DNPM paleontological collection, and Fausto Luiz de Souza Cunha, paleontologist at the MN, sent the material collected by Carvalho and Vidal, respectively, to Herculano Alvarenga, as Souza Cunha was aware of his interest in paleornithology (Alvarenga 1983a ).
Alvarenga prepared and restored these fossils and concluded that most belong to a single species, which he described in 1983 as the new genus and species of small opisthodactylid Diogenornis fragilis , honoring paleontologist Diógenes de Almeida Campos for supporting his research. The species was considered a possible ancestor of the living rheas and one of the oldest known ratites (Alvarenga 1983a ).
The fragments collected by Vidal and Carvalho contained yet another species, which Alvarenga briefly alluded to in 1983 and described in 1985 as the new psilopterid genus and species Paleopsilopterus itaboraiensis (Alvarenga 1983b , 1985a ). It was considered the oldest known phorusrhacoid.
In 1997, paleontologists Robert F. Baird (Melbourne University, Australia) and Patricia Vickers-Rich (Monash University, Australia) described the genus and species Eutreptodactylus itaboraiensis from an incomplete tarsometatarsus collected by Vidal in 1950 and deposited in MN, of which the study was allowed by Souza Cunha. It was considered the oldest known cuculid, probably from a new subfamily, and one of the oldest records for an extant family (Baird and Vickers-Rich 1997 ). Unfortunately, the fossil was lost after being studied, leaving only casts and illustrations in collections.
Mourer-Chauviré ( 1999 ) pointed out the similarities between the Paleogene fossil avifaunas of Europe and South America (including the records from Itaboraí and Taubaté), in addition to recent discoveries that indicated that a similar fossil avifauna was also present in North America. It was possible to define an association of terrestrial or non-marine aquatic taxa that constitute a continental basal avifauna common to Europe, North America, and South America, of which some elements survived only in the last. In the face of the inability or low flight ability of some of these components, Mourer-Chauviré proposed these taxa underwent diversification during the Late Cretaceous or the Paleocene, and dispersed through land routes that existed then, from South America to North America and then Europe, or the other way around.
Alvarenga, Gerald Mayr (Forschungsinstitut Senckenberg, Germany), and Julia A. Clarke (University of Texas, USA) described in 2011 several avian remains present in the MN collection, composed of indeterminate taxa (including material perhaps belonging to Eutreptodactylus itaboraiensis ) and the new genus and species Itaboravis elaphrocnemoides (Mayr et al. 2011a ). The associated elements of this newly described taxon, briefly alluded to by Mourer-Chauviré ( 1999 ), show characters akin to those of the European Paleogene genus Elaphrocnemus , associated with the Cariamiformes.
Furthermore, phalanges referable to Diogenornis fragilis and Paleopsilopterus itaboraiensis and other indeterminate bones were reported in subsequent publications (Taranto and Bergqvist 2009 , 2010 , Taranto et al. 2009a , Alvarenga et al. 2011 , Metello et al. 2012a , 2012b , 2014 , Metello 2013 , Metello and Bergqvist 2014 ).
1.3.3. Concluding remarks
As observed in the Paleocene of Europe, the Itaboraí paleoavifauna seems to include a significant number of representatives with little or no flight capacity. Whether this indicates particular characteristics of early Cenozoic ecosystems, such as the absence of large predatory mammals, remains to be addressed in future studies (Mayr et al. 2011a ). However, large terrestrial predatory crocodilians were recorded in the basin (Pinheiro et al. 2018 ). The associated paleofauna includes marsupials, litopterns, notoungulates, astrapotheres, xenungulates, and cingulates (the site is regarded as the “cradle of mammals” in Brazil), chelonians, lizards, snakes, crocodilians, anurans, caecilians, and gastropods. Fish or any benthic animals were not found, possibly because the lake that filled the original tectonic depression was formed by scalding thermal waters and/or due to the high concentration of calcium carbonate dissolved in its waters (Bergqvist et al. 2006 , 2009 ). Fossil plants and microfossils of ostracods and palynomorphs were also found (Bergqvist et al. 2009 ). Moderately dense woodlands or forests existed in and around the basin area, in a hot and humid tropical environment (Paula Couto 1970 , Alves 2012a , Woodburne et al. 2014a ).
With the cessation of extractive activities on the site in 1984, water drainage was also interrupted, which started to accumulate in the about 70-m-deep depression left by excavations, forming a lake currently used for the water supply of the São José neighborhood. This prevents new geological studies and collection activities (Bergqvist et al. 2006 , 2009 , Kellner and Campos 1999 ).
On 2 September 2018, a large fire heavily damaged the MN, severely affecting its paleontological collections. The status of its important avian materials (encompassing fossils from the São José de Itaboraí Basin, Tremembé Formation, and various quaternary paleontological and archeological sites) deposited there is still unknown, as the recovered remains are still being inventoried (L.B. Carvalho, personal communication). Some fossils were likely saved by being in other institutions at the time, such as the remains of Brasilogyps faustoi and Chaunoides antiquus (recent, new pictures published in Carmo et al. 2024 ).
1.4. Tremembé Formation (SP)
1.4.1. General information
Discovered in the first half of the nineteenth century, the Taubaté Basin is a valley approximately 150 km long (northeast-southwest) and width ranging between 10 and 25 km, surrounded by the Serra do Mar and Serra da Mantiqueira mountain ranges, between the municipalities of Cachoeira Paulista and Jacareí in Eastern São Paulo (Melo et al. 2007 , Kellner and Campos 1999 , Couto Ribeiro 2010 ). It is divided into the Pindamonhangaba Formation and the Taubaté Group, the latter divided into the Resende, Tremembé, and São Paulo Formations (Riccomini 1989 ).
The Tremembé Formation ( Fig. 1 .7) is of lacustrine origin, with an alternating sequence of pyrobituminous shales and bentonite clays, and is generally attributed to the late Oligocene or the early Miocene (late Deseadan SALMA, 24–22 million years BP—Mayr et al. 2011b). The fossil birds of the Formation, comparable to European finds of a well-defined age, support this hypothesis (Alvarenga et al. 2007 ). It is the most fossiliferous unit of the Taubaté Basin, with major contributions to Brazilian paleontology, whether in paleobotany or paleozoology of invertebrates and vertebrates (Alvarenga 1988 ). Particularly crucial for paleornithology, this is the locality with the most avian paleospecies described in the country.
Its main outcrops are in Taubaté, Tremembé, and Pindamonhangaba, being commercially exploited for montmorillonite clay (Couto Ribeiro 2010 ). Most fossils come from Fazenda Santa Fé, belonging to the Sociedade Extrativa Santa Fé, which corresponds to a smectitic clay exploration quarry, located in the Padre Eterno neighborhood, near the right bank of the Paraíba do Sul River, in Tremembé. Its fine sediments preserve, under different fossilization processes, a wide variety of taxa, from microfossils, ichnofossils, and remains of invertebrates, vertebrates, and plants, making it one of the most important geological and paleontological sites of the Brazilian Cenozoic. Despite its importance, much of its information is being lost with the progressive exploration of the quarry (Bernardes-de-Oliveira et al. 2002 ), although, paradoxically, without this process, the systematic paleontological exploration in this site would be practically unfeasible (Couto Ribeiro 2010 ).
Fossils from the Formation were described as early as 1898 in a study addressing fish remains (Paula Couto 1958a ). In 1915, the German-Brazilian zoologist Hermann von Ihering (1850–1930), then director of the Museu Paulista, sent a fossil feather preserved as a carbonized trace (Davis and Briggs 1995 ) from the Taubaté Basin to be examined by the American paleontologist Robert W. Shufeldt (1850–1934). Shufeldt ( 1916 ) published the description of the specimen, associating it with a large species, and noted it was of particular interest as it furnished evidence that “highly developed birds” existed in that Formation.
Another feather from the shale levels was described by the paleoichthyologist Rubens da Silva Santos (1918–1996) in 1950 as belonging to a passeriform. Gilberto H. William found the fossil in July 1948, while collecting along with Silva Santos and Alberto Lopa at the now-deactivated Mina Nossa Senhora da Guia during a paleontological survey carried out by DGM of DNPM. This mine is currently deactivated (Alvarenga 1988 ).
Most of the avian bones found in the Formation were described by the paleontologist Herculano Alvarenga (see below). Besides this, several other non-osteological vestiges of birds in the deposit were described. Prado and Anelli ( 2014a ) first reported two additional feathers (a contour feather and a rectrix) from the gray shales of the Sociedade Extrativa Santa Fé quarry. Prado and Anelli ( 2015 ) performed Scanning Electron Microscopy (SEM) analysis on the contour feather finding microbodies interpreted as eumelanosomes, suggesting the feather was dark colored. Prado et al. ( 2016b ) described them as preserved carbonized traces (although they show different degrees of preservation due to taphonomic processes) and tentatively attributed them to aquatic taxa. Prado ( 2017 ) analyzed them geochemically and microscopically along with specimens from the Crato Formation.
Castro et al. ( 1988a , 1988b ) reported coprolitic nodules associated with birds from Fazenda Santa Fé. It was possible to infer data on the ecology of these animals, demonstrating they had varied eating habits, in addition to information on the evolution of the Tremembé Formation’s sedimentation paleoenvironments. Another coprolite occurrence was depicted later by Souto ( 2012 ). Carmo et al. ( 2023 ) reported 20 specimens (under 19 collection codes) from Fazenda Santa Fé collected in October 2018, and their study constitute the first paleoparasitological analysis of avian coprolites for this Formation. Four of them tested positive for helminth eggs. Three morphotypes of Ascaridina encompassing 16 eggs, an isolated egg referred to the Trichocephalida, two eggs of Spirurina, and an egg of another nematode morphotype were identified. Macro and microstructure analysis revealed the presence of fragments of bony fish and plants, as well as arthropod appendages, which indicates the producers of the coprolites were omnivorous.
Garcia ( 1993 ) mentioned feather impressions while briefly reporting the fossils of the Taubaté Basin in the UNG collection (Garcia and Saad 1996 ), from the Fazenda Santa Fé, Mineração Aligra S/A, and Nossa Senhora da Guia deposits in Tremembé and Taubaté. Additionally, she reported fragments of eggshells, but without attributing them to any vertebrate group. Azevedo and Carvalho ( 1998 ) reported small and thin eggshell fragments from the upper level (dark shales) that presented a structural arrangement strongly reminiscent of the ratite pattern. These occurrences were the first (and only, so far) records of eggshells for this unit, and Azevedo and Carvalho’s record was the first confirmed fossil avian eggs for the country.
A footprint tentatively assigned to Gruipeda isp. was reported by Lima et al. ( 2023 ) from the shales of Sociedade Extrativa Santa Fé. The small tetradactyl trace was likely produced by a member of Opisthocomidae, Rallidae, or Quercymegapodiidae. This is the first assertive record of this nature for the country.
1.4.2. Alvarenga’s studies on fossil birds
After the first studies done mainly by Danish researchers in the nineteenth century, the next big step, which would effectively be a new beginning in the study of fossil birds in Brazil, would come with the work of Herculano Alvarenga on the Taubaté and São José de Itaboraí Basins (Alvarenga 1992 ). His studies and influence on the subject essentially started the age of modern research of this nature in the country, and this could easily earn him the title of father of Brazilian paleornithology.
Herculano Marcos Ferraz de Alvarenga ( Fig. 4 B) was born in Taubaté in 1947 and, from an early age, showed an avid interest in birds, gathering a collection of skins he prepared himself. He graduated from the Faculdade de Medicina de Taubaté in 1973, specializing in orthopedics. Two years later, he joined the institution as a professor, and, in 1977, due to a long college strike, he toured the Taubaté Basin in search of fossils, where he came across the remains of a huge bird in the clays of Fazenda Santa Fé (Mezzalira 2000 ). After analyzing these remains at his home for a while, he sought the help of geologist Diógenes de Almeida Campos of DNPM. The two exchanged correspondences about the material, and Campos encouraged Alvarenga to continue his studies and publish a scientific article on the bird, starting his paleontological career (Pivetta 2003 , Moon 2012 , Andrade 2017 ). Financial support from the National Geographic Society was received for fieldwork in the Tremembé Formation (Alvarenga 1995b).
The well-preserved fragments of the bird’s skeleton were collected between 1977 and 1978, with the help of local workers (Alvarenga 2008 ). They were found in the bentonite clay some 2 or 3 m below the pyrobituminous shales, scattered in an area of approximately 100 m 2 (Alvarenga 1982 , 2003 ). After restoration, the material provided the incomplete skeleton of a large bird ( Fig. 4 .A), described as the new phorusrhacid species Physornis brasiliensis in 1982, initially classified in the genus originally described for Argentinean material (Alvarenga 1982 ).
Tremembé Formation, the work of Herculano Marcos Ferraz de Alvarenga: first photo of Paraphysornis brasiliensis remains, when their restoration began (1977) (A); Alvarenga with the reconstructed replica skeleton (B); Museu de História Natural de Taubaté (C) (H.M.F. Alvarenga).
In January 1978, at the same locality and level as the phorusrhacid fossils, new fragments were collected, this time of a cathartid. Alvarenga analyzed this material at the LACM, and this study resulted in the 1985 description of the new genus and species Brasilogyps faustoi , the oldest representative of the family in South America and somewhat larger than the living Coragyps atratus (Alvarenga 1985b , 1997 ).
Alvarenga ( 1988 ) described the new genus and species Taubacrex granivora from an incomplete skeleton preserved in a shale fragment Silva Santos sent him. It was found about 6 m deep in 1950 by the Petrobrás engineer Mustafá Hanzagic in Mina Nossa Senhora da Guia, former Petrobrás property in Tremembé (Couto Ribeiro 2010 ). Part of the fossil was lost, probably during collection, but it preserved ossified tendons, feather impressions, and a gizzard containing gastroliths. Alvarenga also reported other fragments of bird bones not yet studied from the same origin (as in Olson and Alvarenga 2002 ).
Between 1984 and 1989, fossils determined as phoenicopteriforms, among others, were collected in the Fazenda Santa Fé. Alvarenga ( 1990 ) described them as the new species Agnopterus sicki and material showing great affinity with the European Palaelodus ambiguus . The study of this material also contributed to the knowledge of the site’s paleoenvironment and geological age.
After comparing the fossils of Physornis brasiliensis with museum specimens in Argentina, USA, and Europe and finding significant differences, Alvarenga (1993) proposed to classify it in the new genus Paraphysornis . This fossil’s importance in the development of paleornithology in the country is indisputable, and its solid and curious figure would make it a symbol of the Brazilian paleoavifauna.
Alvarenga, Silva Santos, and other professors of UERJ found, in July 1989, in the shales of Fazenda Santa Fé, the incomplete skeleton of a small galliform, soon mentioned and depicted by Alvarenga ( 1993b ). Alvarenga ( 1995b ) described it as the new genus and species Ameripodius silvasantosi . It was included in the Quercymegapodiidae, initially described for the European Paleogene, showing to be of particular paleobiogeographic interest, especially in the observed similarities between the paleoavifaunas of South America and Europe (Mourer-Chauviré 1999 ). Eventually, Taubacrex granivora , initially associated with the Rallidae, was transferred to the Quercymegapodiidae by Mourer-Chauviré ( 2000 ). These are the oldest known galliform records for South America (Agnolín 2016b ).
In 1995, Alvarenga joined the doctoral program in zoology at the Instituto de Biociências da Universidade de São Paulo (IB/USP), while still teaching and attending his orthopedics office (Andrade 2017 ). His thesis, conceived as early as 1988 and completed in 1999 (Alvarenga 1999b ), resulted in the publication of the systematic revision of the Phorusrhacidae, along with his supervisor Elizabeth Höfling (Alvarenga and Höfling 2003 ).
Alvarenga ( 1999a ) described the new anhimid genus and species Chaunoides antiquus from several fragments collected between 1978 and 1993 in the clays of Fazenda Santa Fé. He mentioned remains attributed to the family on an earlier occasion (Alvarenga 1993b ). It was the first anhimid fossil species to be recognized, representing a slightly smaller and more slender bird than the smallest living species, Chauna chavaria . In that same year, Alvarenga was honored in the specific name of the new curimatid fish Plesiocurimata alvarengai by Figueiredo and Costa-Carvalho ( 1999 ), whose material he helped to collect in 1986 in the Fazenda Santa Fé.
The American paleornithologist Storrs L. Olson (1944–2021) (Smithsonian Institution, USA) came to Brazil (on one of at least three occasions; Pivetta 2003 ) and worked with Alvarenga on bone fragments from the clays of Fazenda Santa Fé. This study (Olson and Alvarenga 2002 ) described the new genus and species Taubatornis campbelli , the smallest (although comparable in size to Pelecanus thagus ) and the oldest known teratornithid. The bird was syntopic with the cathartid Brasilogyps faustoi , and the Tremembé Formation conforms to the almost invariable association pattern of the two families whenever teratornithids are found. This discovery supports the hypothesis of the family’s origin in South America.
In 2011, Alvarenga, Gerald Mayr, and Cécile Mourer-Chauviré (Université Lyon 1, France) described the new genus and species Hoazinavis lacustris , the oldest known opisthocomiform, from fossils found in 2008. Additionally, they revised and reclassified the fossils of Namibiavis senutae (early Miocene of Namibia) also as an opisthocomiform, leading to the hypothesis of the dispersion of the group’s representatives from Africa to South America through a transatlantic route in a vegetation raft, being the first known example for birds (Mayr et al. 2011b ).
Furthermore, Alvarenga (in Castro et al. 1988b , Alvarenga 1993b , 1997 ) mentioned undescribed Anatidae, Podicipedidae, and Phalacrocoracidae remains, among others. Ardeid remains were also mentioned by Castro et al. ( 1988b ). This material’s state of conservation prevents a more detailed analysis but, in any case, they add to the rich fossiliferous bird record in this deposit.
1.4.3. Taubaté’s Natural History Museum
Over the years, Alvarenga gathered a large osteological collection at home, exchanging copies of skeletons with Brazilian and foreign researchers and institutions. Argentinean technicians Pablo Puerta and Raul Vacca were in Taubaté for two months in 1988 to assist Alvarenga in the Paraphysornis brasiliensis skeleton’s restoration, with two resin copies being ready for exchange in 1990. A total of 12 were produced and exchanged with institutions from Brazil, Argentina, Japan, the USA, and several European countries, but this practice was ended in 2005 due to customs bureaucracies (Alvarenga in Mourer-Chauviré 1988 , Alvarenga in Mourer-Chauviré 1990, Tuffani 2002 , Alvarenga 2007a , Moon 2012 , Venceslau 2014 ).
His prestigious scientific career led the Taubaté City Hall to consider the creation of a natural history museum in 1995. In 1998, the City Hall destined land for erecting a building, which it would also finance, and it was completed in late 2000. The Museu de História Natural de Taubaté (MHNT) ( Fig. 4 C) was inaugurated in 2004, with Alvarenga as its director, being maintained by the Fundação de Apoio à Ciência e Natureza (FUNAT), which was created for this purpose (Tuffani 2002 , Venceslau 2014 , Andrade 2017 ). Its collection currently harbors about 14 thousand items from all geological periods (Andrade 2017 )—including extinct bird fossils from other countries (e.g., Alvarenga 1990 , Alvarenga and Höfling 2003 ). Its importance for paleornithology is highlighted by its osteological collection of modern birds, the richest in Latin America (MHNT 2018 ), which by 1987 already had 500 complete skeletons and by 1994 had almost 1,000 species (Mourer-Chauviré 1987 , 1994 ). This collection allowed Alvarenga to publish several articles on living birds (e.g., Alvarenga et al. 2002 , Antunes et al. 2007 ) and other vertebrate fossil groups (e.g., Soria and Alvarenga 1989 , Vucetich et al. 1993 ).
Paraphysornis brasiliensis is the symbol of MHNT (Alvarenga 2007a ) and was pivotal to its foundation (Alvarenga 2008 , Moon 2012 ). Its remains were originally deposited in DGM to be scientifically described, but it was agreed that they would be part of the MHNT collection upon its creation and inauguration (G. Couto Ribeiro, personal communication). Its original bones were shown to the public for the first time during the 2009 “Museum Week” (Alvarenga 2009 ). The MHNT also has as its mascot an animated version of the bird, “Fisó”, created by the artist Marcos Sachs (MHNT 2018 ). Academic projects were carried out based on the MHNT collection or copies of its materials, such as the use of P. brasiliensis as the basis for an animatronic robot (Almeida et al. 2005 ) and its reconstruction and digitalization with the use of three-dimensional virtual modeling (Lima et al. 2008 ).
By 2022, Alvarenga left the position of director, and the institution, now under the direction of Graziella do Couto Ribeiro, was renamed Museu de História Natural Doutor Herculano Alvarenga, honoring its founder.
1.4.4. Concluding remarks
The Tremembé Formation is interpreted as an ancient shallow and alkaline lake (Alvarenga 1999a )—it probably received waters from tributary rivers and, without connection to the ocean, reached high salt concentrations, which allowed ostracods and decapods to reach large numbers and become basic food for the fossil flamingos (Alvarenga 1990 ). The paleoclimate was probably tropical or subtropical in the valley’s interior where the lake was located and colder in the higher areas around it (Melo et al. 2007 ). The shales suggest an alternation between wet and dry seasons, with the fossils indicating periodic mortality of many small fishes; this probably attracted predators and scavengers such as the cathartiforms Brasilogyps faustoi and Taubatornis campbelli (Olson and Alvarenga 2002 ). The fauna associated with the Formation includes notoungulates, astrapotheres, pyrotheres, litopterns, cingulates, sparassodonts, caviomorph rodents, bats, chelonians, crocodilians, snakes, anurans, caecilians, bone fishes, insects, arachnids, crustaceans, and mollusks. Fossils of plants, palynomorphs, and other microfossils, such as ostracods, sponge spicules, and scolecodonts, were also found (Bernardes-de-Oliveira 2002 , Couto Ribeiro 2010 , Melo and Bergqvist 2010 ).
1.5. Solimões Formation (AC, AM)
1.5.1. General information
Late Cenozoic sediments are present in several localities in the Western Amazon (west of Manaus), and their deposits originated in a fluvial-tidal wetland system that existed before the final establishment of the Amazon River system (Negri et al. 2010 ). They contain a wide variety of aquatic and terrestrial fossil vertebrates, and the most diverse faunas are found in the upper Solimões Formation of Solimões Basin, which in Brazil outcrops in the states of Acre and Amazonas but is also exposed in eastern Peru and northern Bolivia (Souza-Filho et al. 2020 ). Two sites, Niterói and Talismã, had radioisotopic maximum age of deposition reported in 8.5±0.5 Ma and 10.89±0.13 Ma, respectively, corresponding to the Tortonian stage (Chasicoan/Huayquerian SALMAs) of the late Miocene (Bissaro-Júnior et al. 2019 ).
The Solimões Formation ( Fig. 1 .9) is a succession of claystones and lignites with sandstone beds, and its upper part, as observed in the state of Acre, is dominated by sandstones, and contains few clay intervals. Samples of its vertebrate fauna were collected at dozens of sites found along the banks of rivers and on roads (Negri et al. 2010 ), notably, in the Upper Acre River, on the triple border between Brazil, Bolivia, and Peru, outcrops the Acre Conglomerate Member from which most of the avian remains of this Formation come (Guilherme et al. 2023 ). Between 1989 and 1998, the staff of the Laboratório de Pesquisas Paleontológicas (LPP) of UFAC collected several avian fossils at two sites: Cachoeira do Bandeira (between Brasiléia and Assis Brasil) and Niterói (in Seringal Niterói, Senador Guiomard), on the banks of the Acre River, among more than 5,000 bones of vertebrates recovered and identified in the region. Cachoeira do Bandeira was discovered by the RadamBrasil staff in 1976, while Niterói was discovered in 1987 in a joint expedition by LPP and MPEG (Souza-Filho and Guilherme 2015 ).
1.5.2. Avian remains
Brazilian fossil birds in the region have been mentioned in the literature since then, but always in brief and sometimes confusing notes. Campbell (in Mourer-Chauviré 1989 ) mentioned a giant undescribed anhinga. Rancy et al. ( 1989 ) listed the genus Anhinga . Bocquentin and Souza Filho ( 1990 ) mentioned a phalanx that attested to the occurrence of large birds in the Niterói site. Wall et al. ( 1991 ) mentioned fossil material from this group. Alvarenga ( 1992 ), through communication with Campbell, mentioned fossils of an extinct anhingid and a giant phorusrhacid (likely the same record mentioned by Bocquentin and Souza Filho), collected by the staff of LACM. Noriega ( 1992 ) mentioned a very large anhingid, through communication with Campbell. Alvarenga ( 1993b ) mentioned the genus Macranhinga as known from the bordering region between Acre (Brazil) and Bolivia (probably Cachoeira do Bandeira site), from material collected by UFAC. Latrubesse et al. ( 1997 ) and Bocquentin and Silva ( 1998 ) listed the genus Anhinga and an indeterminate phorusrhacid without attributing them to a specific site. Silva et al. ( 1997 ), Bocquentin and Janoo ( 1997 ), and Melo et al. ( 1998 ) reported a new species of Anhinga from Cachoeira do Bandeira (but without describing it). Finally, Negri and Ferigolo ( 1999 ) listed an indeterminate phorusrhacid. Ramos et al. ( 2001 ) briefly mentioned a fragment of avian ulna from the Torre da Lua outcrop at the margins of the Tarauacá River, near Eirunepé, Amazonas.
Campbell, responsible for the first mentions of Brazilian fossil anhingid remains, described in 1996 the species Anhinga fraileyi , from late Miocene Peruvian material from the Patos (LACM 4611; Negri et al. 2010 ) and Cachoeira do Bandeira (LACM 5158; Negri et al. 2010 ) sites. The Patos site was discovered in the 1970s and explored in 1985 in a joint expedition by LPP and LACM (Souza-Filho and Guilherme 2015 ). As the sites of the Acre Conglomerate are in bordering regions, the discovered material was variously attributed to one country or another in the literature [4] .
Brazilian anhingid material would only be formally described in 2003. The ornithologist Edson Guilherme, who started a collection of living bird skeletons in LPP, located and reidentified the fossils of birds that were “lost” within the lab’s paleovertebrate collection. Because of the significant morphological similarity, some fossils were identified as reptiles and stored on the shelves next to hundreds of other fragments, mostly crocodilians. After reviewing the entire collection, Guilherme contacted Herculano Alvarenga to study the material together (Guilherme 2016 ). They described two new species: Anhinga minuta and Macranhinga ranzii , respectively the smallest and the (then) largest known anhingas, in addition to material determined as Anhinga cf. grandis and Anhinga cf. fraileyi , taxa originally described from the USA and Peru, respectively (Alvarenga and Guilherme 2003 ).
Bandeira et al. ( 2015 ) briefly reported on two specimens from Cachoeira do Bandeira that were collected during the RadamBrasil project. A humerus was associated with Anhinga cf. A. minuta and a tarsometatarsus with Anhinga indet.
Guilherme et al. ( 2020 ) published new specimens tentatively attributed to Macranhinga ranzii representing new skeletal elements from a new locality named Cajueiro at the left margin of the Purus River in Boca do Acre, Amazonas (Loboda et al. 2019 ). Morphological and myological data enabled them to recognize the taxon as a powerful swimmer and diver, that was able to proficiently hunt fishes in deep waters (such as the frequently recovered Paleohoplias assisbrasiliensis and Hoplosternum sp.), and to provide new phylogenetic hypotheses and improvements in the diagnosis of anhingid taxa.
Hsiou et al. ( 2022b ) reported anhingid cranial and post-cranial elements among the material collected during the July 2022 expedition carried out by the staff of USP, UFAC, and ICMBio (Instituto Chico Mendes de Conservação da Biodiversidade) in the Upper Acre River, in the region of the triple boundary between Brazil, Peru, and Bolivia, where seven sites were prospected, including new and traditional localities. This is the first record of avian cranial material from the Solimões Formation.
Guilherme et al. ( 2023 ) described post-cranial material from that expedition collected in the Acre Conglomerate member of the Patos site at the Brazil/Peru border. Except for a fragment of synsacrum, collected in a curve downstream from Patos, all material was collected between July 18 and 26. They associated them with Macranhinga and Anhinga minuta and suggested the possibility that three distinct anhingid taxa ( A. minuta and their Macranhinga sp. indet 1 and 2) lived contemporaneously in that locality exploiting different fundamental niches. The smaller A. minuta would not be able to feed on large fishes or dive into deep water like the species of Macranhinga and, like the other anhingids from the conglomerate level, could have foraged in a fluvial deltaic environment with running water interspersing shallow and deep ecosystems. The authors pointed out that this supports the interpretation that the region of the Acre Conglomerate Member was formed in a high-energy environment more peripheral to the great lake where the largest anhingids of the Solimões Formation lived. The striking differences in size between some taxa could be explained, to some extent, by diet-related niche partitioning. The discovery of this material also has implications for understanding the material described as Anhinga fraileyi by Campbell ( 1996 ), which encompassed bones from Patos and Cachoeira do Bandeira localities. Guilherme et al. noted that this is an underestimation of the anhingid diversity from that context, and it corroborates Diederle’s ( 2017 ) idea that some of the fossils attributed to A. fraileyi belong to another anhingid taxon.
Furthermore, Souza-Filho and Guilherme ( 2015 ) mentioned Anhingidae material from the Talismã site on the right bank of the Purus River, Amazonas, and avian material from the Morro do Careca site, in the BR-364 highway, between the municipalities of Feijó and Tarauacá, Acre. Muniz et al. ( 2018 ) reported an indeterminate avian tarsometatarsus fragment from the Talismã site collected by the staff of UFAC and USP in 2015–2016. Diederle ( 2015 , 2017 ) revised the anhingid fossil species from South America, treating as Macranhinga sp. the specimens that Alvarenga and Guilherme ( 2003 ) determined as Anhinga cf. grandis and Anhinga cf. fraileyi , and published (Diederle 2016 ) a contribution on the body mass and means of locomotion of Anhinga minuta . Kachniasz and Silva-Caminha ( 2013 ) mentioned Anhinga fraileyi from the Patos locality in Acre, likely referring to Campbell ( 1996 ). The phorusrhacid material still awaits a formal description and was briefly summarized by Agnolín ( 2009b ).
1.5.3. Concluding remarks
The associated fauna of the southwestern Amazon region, one of the last before the Great American Biotic Interchange, encompasses a great diversity of crocodilians and chelonians, as well as lizards, snakes, marsupials, caviomorph rodents, ground sloths, cingulates, litopterns, notoungulates, astrapotheres, sirenians, cetaceans, bats, primates, bony and cartilaginous fishes, crustaceans, and mollusks (Latrubesse et al. 1997 , 2010 , Bocquentin and Silva 1998 , Alvarenga and Guilherme 2003 , Cozzuol 2006 , Bocquentin and Melo 2006 , Negri et al. 2010 ). The anhingids are good indicators of what the environment was like in the region, being attributed to marginal forests of clear lakes and slow-flowing rivers, and along with the associated fauna and flora, indicate an area with large water bodies surrounded by open vegetation mixed with gallery forests that were subject to water fluctuation levels in a tropical seasonal to tropical wet-dry climate (Alvarenga and Guilherme 2003 , Latrubesse et al. 2007 , Negri et al. 2010 ).
The extreme deep diving capacity presented by this group was probably one of the factors that drove its evolutionary history and its extinction (Diederle and Agnolín 2017 ). The radiation of these large anhingids in South America appeared during the Miocene, in tropical and subtropical climates, and disappeared at the beginning of the Pliocene (the Pliocene/Pleistocene age attributed to Giganhinga kiyuensis is disputed), probably due to the combination of deteriorating climatic conditions, the regression of the epicontinental seas and the subsequent disappearance of several freshwater environments, the arrival of carnivorous placental mammals from North America, and the probable competition with the phalacrocoracids, better adapted to survive in more severe climatic conditions (Cenizo and Agnolín 2010 ).
1.6. Lagoa Santa Karst (MG)
1.6.1. General information
Being an object of study since the early nineteenth century, the Lagoa Santa region ( Fig. 1 .10), in the state of Minas Gerais, is the best-known and most researched karstic landscape in Brazil, where it has been one of the first places to be the object of systematic paleontological and archeological research. It is about 30 km north of Belo Horizonte, comprising nine municipalities, in total or part, in the São Francisco sedimentary basin, where the rocks of the Bambuí Group are deposited. It bears the highest density of caves in Brazil, with nearly 1,400 known, many of which provided a wealth of prehistoric material. The fossil finds, with age ranging from the middle Pleistocene to the early Holocene (360,000–9,100 years BP), along with the archeological material, sparked a wide range of interdisciplinary scientific studies, from physical to biological and socio-cultural fields, and are especially important to the question regarding the antiquity of human occupation in the Americas. Albeit unique in some respects, this karst shares similarities with other sites in limestone areas throughout the states of Minas Gerais, Bahia, and Goiás (Auler 2020a , 2020b , Galvão and Peñaranda 2020 ).
Although the proximity to Belo Horizonte (the new State capital established in 1897) made it easily accessible to generations of researchers, it also brought a wide array of threats to the karst’s integrity, some already present since colonial times. The marked increase of human occupation in the last several decades, along with mining and industrial processing, results in degradation through landform and cave destruction, impoverished ecology, air, soil, and water pollution, and damage to paleontological and archeological sites. Since 1980, several protected areas of varying usage categories have been implemented in the karst territory, from federal, state, and municipal initiatives, such as the Lagoa Santa Karst Environmental Protection Area (established in 1990). However, they are still incomplete and fragile and face several challenges regarding applying their concepts and assumptions (Alt and Moura 2020 ).
The karst is intensely researched to this day, and much of the early international attention it received was due to the studies of the Danish naturalist Peter Wilhelm Lund (1801–1880) ( Fig. 5 A). Although not all caves visited by Lund in the early years of his research were part of what today is referred to as Lagoa Santa Karst (such as Gruta de Maquiné), these reports are a necessary inclusion in any study regarding him or the Lagoa Santa region due to their essential contribution to Lund’s initial trajectory (Auler and Piló 2016 ), and, therefore, a requisite to the history of Brazilian paleornithology.
1.6.2. Lund’s studies on fossil birds
Paleontological studies began in Brazil with natural history expeditions carried out during the nineteenth century by foreign researchers, with fossils discovered in caves in Minas Gerais as early as 1809 (Cartelle 2012 ). In this context, during his second and permanent visit to Brazil, Lund established himself in the town of Lagoa Santa, after a fortuitous encounter with his fellow countryman Peter Claussen (1807?–1860?—Holten and Sterll 2011 ) in the town of Curvelo. Claussen showed Lund several of the saltpeter-bearing caves in the region, which he commercially exploited and sold the fossils he found to European museums [5] . This shifted Lund’s focus from general botany and zoology to paleontology, immediately putting him on a path of notable specialization that distinguished him from various other naturalists who explored Brazil during that century (Paula Couto 1950 , Cartelle 1994 , Auler and Piló 2016 ).
Lagoa Santa Karst, the Danish researchers responsible for the first studies on Brazilian fossil birds: Peter Wilhelm Lund (1801–1880) (A); Johannes Theodor Reinhardt (1816–1882) (B); the only known picture we can attribute to Gustav Oluf Bang Winge (1855–1889) (C) (Det Kongelige Bibliotek).
Between 1835 and 1844 (Padberg Drenkpol 1927 ), Lund inspected over 800 caves in the region with the aid of the locals (Lund 1845 –1849), although only a small number contained fossils. He published, besides paleontology, essential contributions to archeology, karst geomorphology, and speleology (Auler 2020a ).
Although Lund focused on studying fossil mammals, he was responsible for the first published records of fossil birds in the country. In his first thesis about his research in the region, written in 1836 and published the following year (Lund 1837 ), he described the cave Lapa Nova de Maquiné (now Gruta de Maquiné, located in the current municipality of Cordisburgo) and mentioned bones of various animals, including birds, in the stalagmite crust that lines the floor of one of its chambers. In the same study, when discussing the remains found in that cave, he mentioned bones of a pigeon-sized bird found next to the remains of a giant sloth, located underneath the stalagmite crust.
Lund also reported traces of owls in Lapa Nova de Maquiné. He associated with predation by Strix perlata (= Tyto furcata ) the large number of rodent and bat bones found in the stalagmite crust and the cavities of the stalagmite masses rising above it, in various states of preservation. He also mentioned bones of this species of owl in several places in the cave, mainly in the area dubbed “Castelo de Fadas”, in its sixth chamber and the passage of access from the fifth to the sixth room, deposited freely on the stalagmite layer and showing strong incrustation, or in the crust itself.
In his first thesis on the ancient Brazilian fauna, written in 1837 and published in 1841, Lund ( 1841a ), when mentioning the first cave he studied, near the village of Cachoeira do Campo, noted he found mounds of remains of small mammals and birds (the latter making up approximately 20 of every thousand individuals), of varying ages, associating their origin with the owls. He even kept some live specimens in his house to study the formation of pellets of undigested material they regurgitate, of which the most resistant fragments make up the remains found in the caves. Remains of the owls’ feeding activities in the caves were mentioned by Lund several times in further works (Lund 1839 , 1841b , 1841c , 1846 ).
In the introduction of his third thesis on the ancient Brazilian fauna, written in 1838 and published in 1841, Lund ( 1841c ) stated that, shortly after his previous thesis, he intended to publish a preliminary report on the vast collection of bird material he found; however, the numerous new mammalian remains made him change plans to complete this already ongoing project. In the same work, to understand why bats are rarer in older deposits than in recent ones of the region, Lund speculated on the nature of the predatory birds responsible for these remains, since the fossil bones also had no signs of crushing and, therefore, could not be the result of the action of predatory mammals. Through the differences he noticed between the composition of the fossil remains—the occurrence of larger animals (e.g., armadillos), reptiles, more birds, and fewer bats—and the modern ones, he speculated the predatory birds might have been a species of larger size or strength. He noted that as “owls do not feed on reptiles”, the predators would probably have been diurnal raptors. This hypothesis was also included in Lund ( 1839 ) and was commented on by Valle and Carnevalli ( 1973 ).
In an appendix to his observations on Brazilian fossil animals, written in 1840 and published in 1842, Lund mentioned the discovery of numerous avian remains, including “two species of rheas, one of which is much larger than the living species” (Lund 1842b: 130). This communication was also included in Lund ( 1840 ).
In his fourth thesis on cave fossils, written in 1841 and published in 1842, Lund ( 1842c ), having sent in the previous year the last supplement of his studies on the fossil mammals of the region (see Lund 1842b ), revealed he had planned the elaboration of a work on the fossil bird finds during the first rainy season, of which he had a large number and had not yet fully examined. However, due to new information he obtained on the fossil mammals, he again occupied himself with them. Nevertheless, he sent a small treatise on the bird remains he had found attached to this thesis. This treatise would be published later in Selskabets Skrifter , according to the editorial staff of Det Kongelige Danske Videnskabernes Selskabs Naturvidenskabelige og Mathematiske Afhandlinger (Lund 1842c ). In this fourth thesis, he also mentioned having found human bones associated with the remains of mammals, birds, reptiles, and fishes in Lapa do Sumidouro, part of the current municipality of Pedro Leopoldo.
Despite the aforementioned note, the treatise [6] was not published. According to Winge ( 1887 ), Lund himself decided this, publishing only an extract (Lund 1841d , 1842a ). In this extract, he reported his provisional results, with 34 species [7] belonging to 26 genera: one bird of prey, eighteen songbirds, six climbers, four gallinaceous birds, and five waders, almost all of genera still existing in the region, including Anabates , Dendrocalaptes [sic], Opetiorrhynchus [sic], Crypturus , and Rhea . He also determined some to the species level, all of which correspond to extant ones, such as Cypselus collaris , Anabates poliocephalus , Capito melanotis , Coccyzus cajanus [sic], Perdix dentata , and Crex minuta[8] . From the extinct fauna, he reported a species of the “family” Alectorides [9] , similar in size to the rheas. He concluded by stating that the most important result of this research is the continuity of the rules [10] established from the comparative study between living and extinct mammals, which also applies to the birds (Lund 1841d , Reinhardt 1881 , Paula Couto 1950 ).
Lund ( 1846 ) reported bones of birds, amphibians, reptiles, and mammals from Lapa da Escrivânia V (Winge 1887 , Paula Couto 1950 ), of the Escrivânia complex, in the current municipality of Prudente de Morais. The excavation took place in 1844 and the site provided an estimated total of 7,569,650 individuals (the vast majority small mammals), of which about 350,000 were birds. The remains of birds in this cave were far more numerous than any other and increased his collection numbers for this group tenfold, with a great variety of species, including some of large size. Of the 126 taxa listed by Winge ( 1887 ), only 20 had no representatives in this cave. The bones were lying on loose soil and were not much altered in general; some appeared rather fresh, and others were slightly rolled (Winge 1887 ). The material varies in age, since the filling of the cave, which is large and deep, must have taken time—Winge noted that sorting out the older forms from the newer ones was not possible, but the most notable specimens may belong to the older layers. Lund associated the large number of small bones in a single site with predation by owls, with the larger bones coming from animals that fell into the caves. He estimated as 5,000 years the smallest time interval for the accumulation of these smaller bones, if a couple of owls brought daily, under an optimistic perspective, four prey items, and considering the time between the replacement of the owl couples. This constitutes one of the first qualified attempts to estimate the age of paleontological material with accuracy through empirical observations (Holten and Sterll 2011 ).
While Lund continued to work in the aforementioned cave, another important one was located by two men who made habitual reconnaissance work, Lapa da Escrivânia XI (Winge 1887 , Paula Couto 1950 ), among several others that did not present remains or presented them in amounts that did not offer prospects of rewarding finds (Lund 1846 ). Mammals, reptiles, and birds were found in this cave, and, of the last ones, the bones were much less numerous than those of Lapa da Escrivânia V, with a less significant number of small parts. According to Winge ( 1887 ), most of these bones had a slightly “burnt” appearance, but not much more altered than those of Lapa da Escrivânia V. The work in the Escrivânia complex was Lund’s last cave study, as he thought he had depleted the region’s fossil novelties (Holten and Sterll 2011 ).
In a letter dated 1844 and sent to Carl Christian Rafn, secretary of the Royal Society of Northern Antiquaries, Lund ( 1845 –1849) described Lapa do Sumidouro and its human and other animal bones. This was Lund’s last work on Brazilian caves to be published (Holten and Sterll 2011 ). Regarding the bird material, he reported that it belongs to several species, with a state of preservation “more or less petrified”, deposited on the loose soil or the bottom of the lake inside the cave. Winge ( 1887 ) described these bones as occurring in considerable numbers, of intense brown or reddish-brown color, most heavy, and some rolled.
Among the less remarkable sites regarding species number, Lapa do Capão Seco provided a considerable number of fossils, but most were poorly preserved, fragmented in many pieces, and partially covered with incrustations, which made determination difficult (Winge 1887 ). Other sites where Lund found bird remains include the Lapas da Anta, da Cerca Grande, do Baú, do Marinho II, da Pedra dos Índios, do Periperi I, do Taquaral III, dos Tatus, do Tiú, Vermelha (of Lagoa Santa), da Escrivânia I, III, and IX, da Anna Felicia, dos Coxos, dos Ossinhos, and da Serra das Abelhas (Winge 1887 ). Winge noted that, in general, few bones appear to have undergone more intense alterations than those of the last four sites, of which the remains appear to be somewhat older than those of Lapa da Escrivânia V, but it is not certain how much older they are. This material includes bones of Rhea , the larger Gyparchus , and Crax , which have been “crushed and cemented again”, but have the same appearance of less preserved bones. Remains of other notable forms, such as Chenalopex pugil , do not have a peculiar aspect and are similar to the other remains from their respective caves.
Lund interrupted his work in the caves in 1844 and the components of his collection, with some exceptions, were sent to Copenhagen between 1845 and 1849 (Padberg Drenkpol 1927 , Cartelle 1994 ). The collection consists of more than 100,000 bones, as well as more than 2,000,000 small bones from owl pellets and about 1,300 breccia samples (Seersholm et al. 2021 ). They were donated to King Christian VIII and Denmark, and the reimbursement of Lund’s expenses was used for the collection’s benefit, such as the payment of a curator, and for that position Lund indicated Johannes Theodor Reinhardt (1816–1882) ( Fig. 5 B), his professor’s son. Reinhardt joined the Galathea expedition that left Copenhagen in June 1845 to take care of the shipping of the boxes ( Fig. 6 A) waiting in Rio de Janeiro and visit Lund in Lagoa Santa to better understand the conditions related to the caves, excavations, and fossil fauna (Holten and Sterll 2011 ).
Several changes in Denmark, such as the death of the King, who had a keen interest in natural history, the establishment of democracy (which caused the royal collections to be moved to other institutions), and the violent damage caused by war, greatly affected the sciences and museum system in the country. This prompted a great delay in the proper exhibition, and, consequently, the study of Lund’s collection in the Kongelige Naturhistoriske Museum in Stormgade, which lacked space. The material remained packed and Reinhardt, unable to exercise the function to which he was designated, returned frustrated to Lagoa Santa on two occasions, from 1852 to 1854 and 1856 to 1858 (Holten and Sterll 2011 ).
The collection was finally exhibited adequately in the Kongelige Naturhistoriske Museum by 1860. The new Zoologisk Museum building opened in Krystalgade in 1870, and Lund’s collection remained there for the next 100 years. When this building also became too small, the institution was moved to the new building at the Universitetsparken and the collection returned to the depot but was now conserved in an accessible manner for researchers (Holten and Sterll 2011 ).
Lund identified part of the bird remains in the collection, but he discovered most of the specimens only after finishing his treatise, of which only the extract was published. Of this larger part, only a small portion was mentioned in his catalog. Several were only referred to as “bird bones”, many were still mixed with bones of other animals or had no indication of which cave they were found in, and some were lost (Reinhardt 1881 , Winge 1887 , Paula Couto 1950 ). The lack of recorded information on this material indicates it is unlikely that Lund went into any further investigation about it (Reinhardt 1881 ). Lund’s determinations (reproduced in Winge 1887 ) are in general accurate, but the exceptions are likely the result of his mammal-focused work, as well as of the limited comparative material. Today, the ZMUC maintains two slightly different versions of Lund’s catalog ( Fig. 6 B), both apparently written copies of the original (Hansen 2012 ).
Lagoa Santa Karst: one of the boxes used to transport the fossil material from Brazil to Denmark (A); example of fossil birds (cathartids) recorded in a copy of Lund’s catalog held in the ZMUC (B); drawings of the purportedly alectorid remains, fragment of tarsometatarsus (C) and incomplete pedal phalanx in lateral (D) and front (E) views; (A–B: K.L. Hansen; C–E: after Reinhardt 1881 ).
1.6.3. Other contemporary studies
Other authors contemporary to Lund published about fossil birds from the Lagoa Santa region, but most were simple mentions or reproductions of his studies. Claussen ( 1841 ) described the geological aspects and the fossils found in the region in a volume written to accompany the fossils and mineral samples he sold to European museums (Holten and Sterll 2011 ). He noted that, together with Lund, he visited several caves, of which he said they found “101 species of mammals, 31 species of birds, reptiles, snakes, etc.” (Claussen 1841: 340). Following the list of mammals, which makes up the main part of the material he said they discovered until January 1840, he complemented it with a note stating they found “a large number of bird species, including two species of ostrich [rhea], where one is much larger than the living one”, in addition to reptiles, amphibians, and invertebrates (Claussen 1841: 341). He also mentioned Strix perlata and its activities. These data were almost exact copies of Lund’s observations and, even where they conducted excavations together (such as in Lapa Nova de Maquiné), Claussen only took part in collecting the material, while Lund was the work’s intellectual manager. The publication of this information, threatening Lund’s reputation, worsened their relationship, which was tense most of the time (Holten and Sterll 2011 ). Lund demanded a retraction from Claussen, which did not occur. This led him to send a rectification (Lund 1843 ) to Heinrich Georg Bronn, editor of the reputable German geological journal Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde (Holten and Sterll 2011 ).
Gervais ( 1844b , summary in 1844a ) referred to Lund’s data reproduced in Leonhard and Bronn ( 1843: 236–237) and mentioned the bones of the rhea that is “larger than the two [sic] living species”, aside from adding information about specimens supposedly found by Claussen [11] . He reported that Claussen, who had returned from Brazil, informed him that he had found numerous bird fossils but had no time to sufficiently identify them. However, it was possible to recognize a Cathartes larger than the living species, a Strix , a Caprimulgus , a genus close to Dicholophus , and a Psittacus “or parrot”. Reinhardt ( 1881 ), who had access only to Gervais’ summary (where he only informs that Claussen’s collection allows adding these five records to Lund’s list), reported that this material was discovered in a collection of bones from caves of Brazil that the Muséum d’Histoire Naturelle in Paris had then purchased from Claussen, and that some of these five forms may have been discovered by Lund without Gervais realizing it (see above). He further noted that, if these determinations were correct, Cathartes and Strix are indeed new, but he did not observe any Cathartes or Catharistes larger than the living species in Lund’s collection. However, among the material he collected in the final years of his activities, there are the jaw and other bones of a Gypargus very close to or identical to the living G. papa (= Sarcoramphus papa (Linnaeus)), and perhaps this is the bird Gervais referred to. Winge ( 1887 ), citing Reinhardt for the Cathartes , noted that Caprimulgus and Psittacus are found among Lund’s material. The material classified as Caprimulgus in Lund’s catalog is referred to as G. sp. indet., forte Hydropsalis forcipata by Winge. Psittacus may be cf. Amazona sp., where a humerus fragment was referred to by Lund in his catalog as Psittacus aff. guyanensi (Winge listed it as Chrysotis sp. e minor., vel g. aff. and noted that the term used by Lund is synonymous with Conurus pavua ), or Paraclaravis geoffroyi , of which a humerus was referred to as Psittacus in the same catalog. The genus close to Dicholophus (= Cariama ), which Winge stated he was unable to identify, may refer to Lund’s giant alectorid (Gervais mentioned, as already noted, besides the alectorid, the “species of rhea larger than the two living ones”, which most probably refer to the same material). Since Gervais’, nothing seems to have been published about the material housed in the Muséum d’Histoire Naturelle.
Helmreichen ( 1846 ) was among the many travelers who had been to Lagoa Santa and was received by Lund on two occasions in 1841 (Renger 2002 ). He mentioned that Lund had found more than 100 species of extinct mammals and more than 30 species of vanished birds and reptiles inside the caves.
Liais ( 1872 ) also visited Lund and found the sternum of a hummingbird in a cave in the region. He mentioned it along with information based on the work of Gervais (from which Liais attributed the alleged findings of Claussen to Lund).
Wallace ( 1876 ) mentioned the 34 birds from Lund’s treatise extract and highlighted the “two species of Rhea ”, as well as the large Cathartes and the Cariama or “new allied genre” from Gervais’ work. The most curious, however, is the inclusion of “a new species of the very isolated South American genus Opisthocomus ” (Wallace 1876: 164). It is likely a mistake, as noted by Newton ( 1881 ) and Reinhardt ( 1881 ). Reinhardt further added that Wallace did not seem to have seen or examined the fossils he referred to, and judging by the way he wrote, his intention was only to briefly reproduce third-party communication since he apparently only consulted Gervais’ summary and the letter Lund sent to the Annales des Sciences Naturelles . Perhaps the genus Opetiorhynchus is the source of the confusion. A new genus for the hoatzin family in Brazilian territory would not be published until 135 years after Wallace’s work (Mayr et al. 2011b ). Lambrecht ( 1933: 759), probably unaware of Wallace’s mistake, still referred to remains of the family among the fossil finds, mentioning “Opisthocomidae, from the Pleistocene of the Brazilian ossiferous caves and Eastern Peru (Winge and Lund)”.
1.6.4. Reinhardt’s and Winge’s studies on fossil birds
Further studies on the bird remains discovered by Lund would be published only after his death. Reinhardt ( 1881 , translated into English in 1882) analyzed and commented on how the bird fossils influenced Lund’s conclusion of equating this group with the mammals as to the rules [10] he established between faunas of the past and present, which supported Cuvier’s theory of catastrophism (Cartelle 1994 ).
Reinhardt considered Lund’s rules to be true to some extent. The bird specimens belong to species typical of the American continent, regardless of the proportion of truly extinct species (just as the mammals found in the same deposits). Moreover, the deposits included species that in the present occur regularly only in other parts of South America, even if occasionally some vagrant specimen appears in the region [12] . Yet he doubted the agreement between the two groups regarding the diversity of sizes observed between the extant and extinct mammalian fauna, from which there were both larger versions of living genera and exclusive giant forms of the past, a feature not observed among the birds, except for the giant alectorid of which Lund believed to have found the fossil remains and that, according to Reinhardt, even if correctly interpreted, would not be sufficient to sustain such a conclusion.
To better understand the giant alectorid and its importance on the matter, Reinhardt carefully examined its remains, including possibly associated fragments and illustrations of lost material ( Fig. 6 C–E), and concluded they belong to the living Rhea americana . This result led him to propose there are no indications that the region’s ancient bird fauna had large extinct representatives, thus eliminating the integral parity between this group and the mammals as to the rules employed by Lund. He stated that “the extinct bird fauna of which the remains are found in the caves of Brazil differs from the existing fauna in a much lesser degree than the mammalian fauna which is buried there” (Reinhardt 1881: 153). He further added that, despite the result of the analysis, he believed there may be in fact extinct forms among the fossils found, but that only through a new and careful investigation of the material one can assess the real proportion between vanished and living species.
A thorough examination of the mammalian (including human) and bird remains of Lund’s collection was published under the collective title “ E Museo Lundii ” between 1887 [13] and 1915. His fellow countryman Herluf Winge (1857–1923), a specialist in mammals, wrote most of the monographs that comprise this work, based on the fossil material and Lund’s scientific documents, published or not (Paula Couto 1950 ). The ornithologist Oluf Winge (1855–1889), Herluf’s brother, wrote the monograph dedicated to the bird remains, published in 1887 , with 54 pages and one plate, drawn by Herluf. An addendum addressing the remains of reptiles, amphibians, and bony fishes was published only in 2012 (Hansen 2012 ).
By invitation of Reinhardt, Winge began studying the bird remains in the boreal summer of 1881, the year in which he obtained the title of “ candidatus magisterii ” in Natural History, and he gained the position of assistant in the Zoologisk Museum in 1883 (Collin 1905 ). Reinhardt died in 1882, and Winge’s studies continued under the auspices of Professor Christian Frederik Lütken.
The comparative analysis was based on Lund’s large collection of Brazilian bird skeletons, as well as other skeletons from the museum and Winge’s collection. It resulted in a list of 126 taxa (plus addenda), several probably including more than one species. The vast majority consists of living forms, many of which can still be found in the vicinity of Lagoa Santa. Regarding the avifauna of the region, Winge noted that only the families Anhimidae, Galbulidae, Cotingidae, Pipridae, Mniotiltidae (=Parulidae), and Coerebidae (=Thraupidae: Coerebinae) do not have identified representatives in the fossils, but material of the last three may exist among the large number of indeterminate passeriform bones.
Of the birds that were found in the caves and are not included in Reinhardt’s ( 1870 ) survey of the avifauna of the “Brazilian Campos” [14] , Winge listed little more than two dozen, none of which is particularly strange to the region, including species that are found in nearby areas or had recent remains found in the caves. Lund and Reinhardt, or other naturalists, did not find them alive perhaps by chance.
Winge described a single new extinct species from numerous remains of three sites, the large anatid Chenalopex pugil (now Neochen pugil ). It was the first species of fossil bird (and dinosaur) to be named for Brazil. He also noted that two other forms, Vanellus aff. cayennensi and Gyparchus aff. papae , are either extinct or only represented alive by specimens of much smaller size, but that are close to species that live now in the region anyway. Along with Neochen pugil , these forms are the only ones considerably larger than their living counterparts. Although part of the not completely defined forms (some of which were not mentioned by him) could represent something atypical, Winge noted that one could not change the conclusion that the fossil avifauna of the region is very similar to the living one. Special attention was given to the search for species of these groups without representatives in Brazil or the American continent among the material, but it was without success.
Winge fell ill with lung tuberculosis, which forced him to interrupt his work before passing away prematurely in 1889 (Collin 1905 , Helms 1932 ). His work is important as a starting point for the thorough study of Brazilian fossil birds but, just like the whole “ E Museo Lundii ”, it was not widely distributed in Brazil and there is no easily accessible translation of its content into Portuguese [15] (most of the volumes have, however, French summaries [16] ). Biographical information on him is also very scant, and the only picture that can be attributed to him ( Fig. 5 C) was discovered by us from a hanging portrait in a photograph of Herluf Winge’s living room present in Det Kongelige Bibliotek collection. A compendium of all the biographical data found is in preparation.
Lund’s fossil bird collection was never thoroughly reviewed after Winge’s study. Olson ( 1985a ) noted that, due to probably limited comparative material, even some of the most positive determinations could not be accepted without contestation: 58 taxa, almost half of the total, show some degree of uncertainty regarding their determination. Winge himself did not exclude the possibility that some of the determined fragments could represent related species of other territories or even extinct forms, as also pointed out by Alvarenga ( 1992 ). Vuilleumier ( 1987 , 1993 ) also indicated the necessity of restudying this material using more abundant, recent osteological material for comparison and with modern study techniques. Winge noted that, given the general similarity between the fossil and the living avifauna, this probably would not essentially affect his results. Nevertheless, these factors do not demerit Winge’s work, who had perceived certain traits that would result in the description of a new genus and species over a century later (see below) (Alvarenga and Olson 2004 ).
1.6.5. Further work on bird remains
The next known new species from the region, the ciconiid Ciconia lydekkeri , was described in a convoluted nomenclatural act from remains Claussen sold to the British Museum (Natural History). British paleontologist Richard Lydekker ( 1891 ) thought the Brazilian material belonged to Palaeociconia australis , a taxon described by Argentinean paleontologist Francisco Pascasio Moreno in 1889 . The feud between Moreno and his fellow countryman and paleontologist Florentino Ameghino (and his brother Carlos) fueled the confusion regarding the correct designation of the material, for which Ameghino ( 1891 ) erected the new genus and species Prociconia lydekkeri (see below). Lydekker also associated remains sold by Claussen with Sarcoramphus papa and the extinct Chenalopex pugil . Further material from the Claussen Collection was associated with the ciconiid by Harrison ( 1975 ). Agnolín ( 2009a ) ultimately revised the ciconiid material, considering the description by Ameghino valid and associating it with the extant genus Ciconia .
Souza Cunha and Magalhães ( 1986 ) reported the rare occurrence of bird remains from the Cerca Grande complex in Matozinhos, excavated in 1956 as part of the “Lagoa Santa” archeological project. From this site, 3,425 vertebrate bones of early Holocene age were excavated and housed in MN. Of this material, only about 2,000 bones were identified; birds, together with amphibians and fish, made up about 2% of this number. The remains belong possibly to psittacids and passeriforms and were found in Cerca Grande I, II, and VI. They were interpreted as having anthropic origin (Paula Couto 1958b, Souza Cunha and Magalhães 1986 ).
Souza Cunha and Guimarães ( 1978 , 1981 –1982), responsible for the studies of the Holocene fauna during the Franco-Brazilian Archeological Mission (1971–1976), briefly described the bird remains of pleistocenic and holocenic age found among this material in Lapa Vermelha IV (“Grand abri de Lapa Vermelha (P.L.)”), then in the municipality of Pedro Leopoldo, now part of the recently created (1995) municipality of Confins (about 20 km from Lagoa Santa) (Prous 2016 ). They mentioned a psittacid maxilla, the tarsometatarsus of a large bird found in the same sediments, and numerous passeriforms represented by elements such as mandibles, maxillae, and various long bones, all poorly preserved, preventing a more accurate determination. Alvarenga ( 1996 , 1998 , 2007 ) associated part of this material with the living species Vultur gryphus and Anodorhynchus glaucus or Anodorhynchus leari . These remains were housed in the MN.
Besides the two taxa described in the nineteenth century, only in the 2000s would new species be published from the region, both condors of opposite sides of a line regarding body size. They are the small Wingegyps cartellei , described by Alvarenga and Olson ( 2004 ) from material collected by Lund along with material from Gruta dos Brejões, Bahia, and the large Pleistovultur nevesi , described by Alvarenga et al. ( 2011 ) from Gruta Cuvieri in Matozinhos. Additionally, Alvarenga and Silveira (in Silveira et al. 2008 ) mentioned the presence of undescribed species of Penelope among the material from caves of Minas Gerais, in the Lagoa Santa region, and of Bahia, discovered by the team led by paleontologist Cástor Cartelle.
From the literature, Nascimento and Silveira ( 2020 ) attributed 161 records from pleistocenic to early holocenic age that were determined to a taxonomic level below Aves in the region. Additionally, many undescribed materials await study (e.g., Souza Cunha and Guimarães 1978 , 1981 –1982). As they purportedly belong to living species, some records were mentioned simply as “birds” in the literature, and their younger age and poor conservation support this reduced interest (e.g., Paula Couto 1958b ). This is particularly true for the remains excavated in archeological sites, of which Walter ( 1958 ) noted they are quite common in camp refuse. These were reported on several occasions (e.g., Veloso 1983 , Veloso and Resende 1992 , Cartelle et al. 1998 , Hubbe 2008 , Perez 2009 , Kipnis et al. 2010a , 2010b , Hubbe et al. 2011 , Mayer 2011 , Mingatos 2017 , Chim 2018 ; see Table 4 ).
Seersholm et al. ( 2021 ) studied ancient DNA from samples of approximately 100 bone fragments from Lund’s collection. The bones consist of owl pellets mixed with some bone fragments of larger animals and were collected in Lapa da Escrivania V. The samples were analyzed with bulk bone metabarcoding and showed a high species diversity, including two bird species ( Falco sp. and a rallid), besides indeterminate neognaths. Avian remains were also detected morphologically in all samples. Their study demonstrates the potential of future genetic studies on Lund’s collection.
1.6.6. Concluding remarks
Establishing a chronology for the avifauna described by Winge is difficult. Cenizo et al. ( 2015 ) followed available data (Dias 2004b , Bueno et al. 2013 ) that estimated an age between 12,000 and 8,000 years BP and suggested it may correspond to the stadial event of Younger Dryas in the post-Late Glacial Maximum. Some remains are of more recent age (Reinhardt 1881 , Winge 1887 ). Winge assumed that bodies of water and forest extension could have been more abundant in the region in the past because of the diversity of anatids (at least six taxa versus the four living in the area now) and the presence of large psittacids and the genera Crax and Xiphocolaptes , respectively, among the material collected by Lund. Nevertheless, he noted that drastic changes in the landscape do not seem to have happened according to the general picture of the bird remains found. Cartelle ( 2020 ) used this interpretation of more significant forest extension inferred from the bird remains to support the occurrence of different mammalian megafauna taxa in the region compared with other pleistocenic intertropical sets in Brazil, which included larger giant sloth and proboscidean species. Pollen assemblages recovered in the region indicate a wetter and cooler climate than today, with a mixture of typical cold-climate elements co-occurring with cerrado and tropical forest taxa during the Pleistocene–Holocene transition (Raczka et al. 2013 , 2018 ).
These avian remains are particularly important due to their taxonomic diversity and historical significance, especially due to the ancient avifauna of Brazil being very poorly known, as is South America’s in general (Olson 1985a , Alvarenga 1997 , Tambussi and Degrange 2013 ). They allow a faithful picture of the bird community of the Pleistocene–Holocene transition in the region, which enables us to understand not only the composition of ancient biotas but also the changes in the environments and their faunas over time. This avifaunal assemblage was also included in studies regarding South American biogeography (e.g., Vuilleumier 1984 , 1985 , 1993 , Olson 1985a ). Besides birds, remains of invertebrates, fishes, amphibians, reptiles, and mammals (including humans) were also found in the region (e.g., Lund 1837 , Paula Couto 1950 , Hansen 2012 ). The mammalian megafauna stands out by being numerous and diverse, including proboscideans, giant sloths, glyptodonts, camelids, equids, litopterns, short-faced bears, and saber-toothed cats (Cartelle 2016 ).
1.7. Further records
Besides the aforementioned ones, other notable sites with material described to the family level or that are of interest for their ancient age will be presented in the following subsections, with further details available in the record accounts. Indeterminate fragments of varying temporal and spatial distribution were reported widely in the literature, and as many sites as possible in which they were found are included in Table 4 . It should be noted that this is not an exhaustive survey of these mentions, which permeates especially the archeological literature, in a vast and decentralized manner.
1.7.1. Entre-Córregos Formation (MG)
The Aiuruoca Basin, located between the northern border of Serra da Mantiqueira and the southern border of Serra de Minduri in Planalto do Alto Rio Grande, southern Minas Gerais is composed of the Entre-Córregos and Pinheirinho Formations, and was first described by Santos ( 1999 ). Its sedimentary packing suggests it was deposited on a terminal lacustrine system (Bedani and Haddad 2002 ).
The Entre-Córregos Formation ( Fig. 1 .6) is fossiliferous and composed of pelite in the form of shales with argillite intercalations. Its remains are characterized by plant megafossils, insects, fishes, anurans, coprolites, and palynomorphs (Bedani and Haddad 2002 , Franco-Delgado and Bernardes-de-Oliveira 2004 ). The palynomorphs indicate an Eocene–Oligocene age for the outcropping section of the Formation (Garcia et al. 2000 ), and the leaf architecture of the plant fossils indicates the region had high temperatures and wet weather (Castro-Fernandes et al. 2013 ).
Among these remains, there are also some fossil feathers. They occur in the medium-gray argillaceous, papyraceous shales in the 1.42 m thick basal layer of the Formation, along with rare plant material, fishes, and insects, and abundant pipid anurans. They were collected around the left margin of the Entre-Córregos creek near the road that connects the municipalities of Minduri, Cruzília, Andrelândia, São Vicente de Minas, and Aiuruoca (Bedani and Haddad 2002 , Franco-Delgado and Bernardes-de-Oliveira 2004 ). Brief mentions exist in the literature (Bedani and Haddad 2002 , Franco-Delgado and Bernardes-de-Oliveira 2004 , Castro-Fernandes et al. 2013 ), but unfortunately, these specimens do not seem to have been described in detail nor depicted anywhere. Nevertheless, they attest to the presence of birds in these deposits and give hope for future finds of osteological material.
1.7.2. Gruta dos Brejões and Toca da Boa Vista (BA)
Paleontologist Cástor Cartelle (PUC Minas, then UCMG) and his team collected numerous quaternary bird bones associated with mammalian fauna in caves in middle east Brazil during the 1980s (Mourer-Chauviré 1988 ). In 1980 and 1984, 2,000 bird bones (Cartelle and Santos 1985 , 1,800 according to Penido et al. 2012 ; about 1,000 by 1983 according to Cartelle 1983b ) were collected in Gruta dos Brejões ( Fig. 1 .11) in Morro do Chapéu, Bahia. The roof collapse at three points along the 11 kilometers of the cave created openings that facilitated access to birds, especially in the area dubbed “Salão das Aves”, where most of the material, belonging to non-cave dwelling families, was found. A picture of avian bones being collected in this cave was published by Cartelle ( 1983b ), who also reported beaks among the well-preserved material. In addition, many small bones interpreted as disaggregated from owl pellets by water infiltration were also found in the same area. By 1983 this material already numbered more than 900 items and was being studied by the UFMG and UCMG ornithology staff (Cartelle 1983a ). Cartelle and Santos ( 1985 ) reported a complete ciconiid skeleton and 33 skulls belonging to nine families. By 1988 this material was taken to Taubaté by Cartelle’s student José Enemir dos Santos to be analyzed in collaboration with Herculano Alvarenga (Mourer-Chauviré 1988 ). By 2012 it was still being revised, and three taxa were determined to the species level, including the extinct condor Wingegyps cartellei (Alvarenga and Olson 2004 , Silva et al. 2012b , Penido et al. 2012 ).
Unregistered and unidentified quaternary bird fossils figure among the MHNJB/UFMG paleontology collection (Greco and Cozzuol 2012 ). From Toca da Boa Vista ( Fig. 1 .12) in Campo Formoso, Bahia, some advance was made regarding the cataloging and determination of material collected in the 1980s by Cartelle and his team (Silva 2010 , not consulted by us; M.A. Cozzuol, personal communication). Silva and Cozzuol ( 2010 ) reported seven taxa associated with dry savannas, of which the preservation status suggested an age between the late Pleistocene and Holocene. Unfortunately, the MHNJB suffered a fire on 15 June 2020, but the paleontological collection was not affected (M.A. Cozzuol, personal communication).
Maia et al. ( 2010 ) pointed out the scientific potential of the paleornithological collection of PUC Minas collected by Cartelle and colleagues and noted it harbors more than 2,000 elements including sterna, coracoids, radii, ulnae, femora, tibiotarsi, tarsometatarsi, and synsacra. Most are well-preserved and not studied and come from various sites in Brazil. The collection was being revised and reorganized at the time.
Furthermore, Leoni et al. ( 2024 ) reported cathartid remains from Toca da Boa Vista. The evidence points out that the bird died inside the cave and was later partially scavenged by a small felid.
1.7.3. São Raimundo Nonato region (PI)
Guérin et al. ( 1993a , 1993b ) briefly reported avian remains of quaternary age from the Toca da Janela da Barra do Antonião rock shelter in Coronel José Dias, São Raimundo Nonato archeological area ( Fig. 1 .13) in southeastern Piauí. It is the most important one of the hundred cavities situated in a reduced karstic area in the southeastern vicinity of the Parque Nacional Serra da Capivara, which provided a wealth of archeological and paleontological remains and paleoclimatic indicators. Discovered in 1986, this large rock shelter was excavated up to 1990 by archeologist Niède Guidon. It further provided a rich pleistocenic fauna, human burial, rock paintings, and lithic tools associated with megafaunal remains (Guérin et al. 2002 ). Cenizo et al. ( 2015 ) suggested an age of 9,670±140 years BP for this avian assemblage based on data by Martin ( 1996b ).
The avian remains come mainly from the upper levels of sector C (0.05–0.50 m deep) and from the surroundings of a human skeleton with an estimated age of about 9,700 years old (0.55 m deep), as well as from slightly deeper levels of sectors A and B (0.74–1.37 m deep). No significant differences were observed between the composition of these two parts of the deposit other than that the upper levels of sector C are richer and more diversified.
Due to the lack of comparative material for the living South American avifauna in European osteological collections, it was not possible for them at that moment to determine the material at the generic and specific levels. Later, with the aid of American collections, Guérin et al. ( 1996 ) preliminarily reported, besides indeterminate Passeriformes, 32 taxa (Simões 2001 ; later reported the total number to be 42). Although minimum numbers of individuals (both adult and young) were provided for most taxa, no skeletal elements and their conservation status were specified, nor where the material was deposited. Guérin et al. ( 1993a ) included, in the preliminary report, the occurrence of Phasianidae among the material, a record which was not present in their other publication of the same year nor in that of 1996.
Most of the recorded taxa can still be found living in the Parque Nacional Serra da Capivara and adjacent areas, with the absence of some possibly indicating environmental changes that caused some reduction of forested areas. There is also a notable scarcity of aquatic forms among the rock shelter material when compared with the living avifauna (two versus 12 families, respectively), possibly due to the alimentary preferences of the ancient human inhabitants or the raptorial birds responsible for the accumulation of the material, as aquatic and semiaquatic mammals and large amphibians are well represented.
More avian materials from levels 1 and 3 of sector A of Toca da Janela da Barra do Antonião were later reported by Moraes ( 2014 ), including apparently burned indeterminate material. She also attributed to owls the pellets of small rodent bones found at the same site.
The surprising absence of Rhea among the remains of Toca da Janela do Antonião was noted by Guérin et al. ( 1993b , 1996 ), a bird which is well represented in the Nordeste Tradition rock paintings dated between 12,000 and 6,000 years BP, though it no longer occurs in the region today. From Toca do Serrote das Moendas, a new site also in the vicinity of Parque Nacional Serra da Capivara that began to be excavated in 2006 under the direction of Elaine Ignacio, Guidon et al. ( 2009b ) first mentioned the discovery of Rhea[17] as part of a rich late Pleistocene fauna associated with human remains and lithic tools. Later, Faure et al. ( 2010b ) associated the fossil with Rhea fossilis , a species first described from Argentina (a reevaluation of the original material considered it to belong to the living Rhea americana ). Remains of Blastocerus and Hydrochoerus were also first unambiguously reported from these deposits. The nature of this publication, in a volume specialized in rock art, prompted this record to be largely ignored in publications concerning the bird fossil diversity in Brazil.
1.7.4. Natural tanks in Itapipoca (CE)
Natural tanks with associated fossiliferous deposits are found throughout Northeast Brazil and primarily preserve late Pleistocene megafauna remains (Ximenes 2009 ). Bird remains are rare in these deposits, with just a handful of records in the literature.
During January and February 1961, an MN expedition led by paleontologist Carlos de Paula Couto with the collaboration of geologist Fausto Luiz de Souza Cunha took place in the João Cativo paleontological site in Itapipoca ( Fig. 1 .14), Ceará. They discovered bird material in a tank deposit (tank 2) and reported it as indeterminate neognath orders and genera in their expedition reports. These are the first records of bird fossils from the Northeast region of the country (Souza Cunha 1961 , Paula Couto 1961 ). Later, part of this material was associated with the genus Rhea (Paula Couto 1962 , 1980 ) and a hawk (Metello and Araújo Júnior 2012 , 2013 ). This accipitrid record suggests an association between forest environments and open areas for the Itapipoca region during the Pleistocene (Metello and Araújo Júnior 2012 , 2013 ). Patusco et al. ( 2016a , 2016b ) briefly described part of this material and other fragments from tanks 2 and 3. Costa et al. ( 2024 ) restricted part of the material to Rhea americana and an accipitrid.
Furthermore, in Itapipoca, another tank deposit in the Jirau paleontological site (tank 1) provided indeterminate bird material during work done between 2003 and 2008 by researchers Celso Lira Ximenes and Antônio Sílvio Teixeira dos Santos (Araújo Júnior 2012 , Araújo Júnior et al. 2013 ).
1.7.5. Archeornithological remains
Avian subfossil (i.e., non-completely fossilized and/or of recent holocenical age) remains in Brazil are generally studied under the scope of zooarcheology, as most discovered material is associated with archeological sites. These remains are common in pre-colonial (i.e., before the arrival of Europeans) sites across the country, be they rock shelters, shell mounds (sambaquis), or cerritos, but are rarely determined to a specific level or depicted (Chahud 2021 ). Zooarcheological research in Brazil has only recently started to develop, and the identification and study of avian remains still fall behind that of mammals and fishes concerning the vertebrates but is nevertheless a promising field, on both prehistoric and historic sites (Rosa 2008b ).
The earliest mention we found of a bird specifically from an archeological site is that of Wiener ( 1876 ), who reported a parrot skull from a sambaqui in the Tavares River in Ilha de Santa Catarina. From this period, Löfgren ( 1893 ) also reported bird remains among the material found in 1884 in the sambaquis of Ilha do Casqueiro and Palmeiras near Santos, besides occurrences from sambaquis in the Mar Pequeno area, all in São Paulo. Meyer ( 1896 ) reported bird bones in a sambaqui in Magalhães, east of Laguna, Santa Catarina, and Krone ( 1914 ) mentioned arrowheads made of bird bones from sambaquis in the valley of Rio Ribeira de Iguape in São Paulo. Mentions of avian remains only started to be more common in the literature from the 1970s onwards, and the first study focusing solely on birds was only published in 2014 (Cardoso et al. 2014 ).
Most of the material was published as indeterminate birds ( Table 4 ) in faunal checklists, due to a combination of poor state of preservation, lack of comparative material, and the focus of zooarcheological studies directed toward mollusks, fishes, and mammals. Thus, much material in collections remains indeterminate and unpublished. The determined material is generally that of species of food interest (e.g., rheids, tinamids, spheniscids, and rallids), providing data on feeding and occupation patterns of ancient human populations and the ecology of ancient environments (Rosa 2008b ), but remains interpreted with ritualistic or non-anthropogenic origins were also described. Much of the archeornithological record is also parallelly present as tools prepared from long bones (such as arrowheads) or ornaments, and those are generally simply referred to as having an avian origin without further taxonomic investigation, in good part due to their modified aspect. Eggshells have also been reported and interpreted as consumed for food or as part of ornaments or tools.
The discovered and published material comes mostly from coastal sites in the South and Southeast regions of the country. In the South region, we can highlight the work done by archeologists associated with the IAP/Unisinos such as Pe. João Alfredo Rohr (1908–1984), Pedro Ignácio Schmitz, André Osório Rosa, and André Luiz Jacobus (1957–2016), mainly in Rio Grande do Sul and Santa Catarina, besides work done in Goiás and Mato Grosso do Sul. In the Southeast region, the project “Saquarema-Rio de Janeiro: Pré-História e Paleoambiente”, coordinated by archeologist Lina Maria Kneip (1938–2002), uncovered important avian material from the Beirada, Moa, and Pontinha sambaquis starting in 1987. There is no systematic study published focusing on avian remains from Brazilian archeological sites, but Thayane Braga de Souza Patusco (MN/UFRJ) is currently working on this subject for her master dissertation (Patusco 2021 , 2023 ).
1.8. Brazilian contribution to international research
Brazilian researchers also contributed to international research on fossil birds. Alvarenga described a flightless bird from the Cretaceous of Argentina ( Patagopteryx deferrariisi ; Alvarenga and Bonaparte 1992 ), an anhingid from the Miocene of Chile ( Meganhinga chilensis ; Wall et al. 1991 , Alvarenga 1995a ), and phorusrhacids from the late Pleistocene of Uruguay (Alvarenga et al. 2010 , Jones et al. 2016 , 2017 ). Riff and Kellner (in Riff et al. 2004 ) published on an avian vertebra from the Cretaceous of Morocco. Silveira (in Grellet-Tinner et al. 2012 ) published on an aquatic twig-nest with phoenicopteriform eggs from the Miocene of Spain. Carvalho (in Agnolín et al. 2017b ) published on a fossil feather from the Jurassic of Kazakhstan. Additionally, Late Cretaceous avian remains were unearthed in Vega Island as part of the Paleoantar project of Programa Antártico Brasileiro (Proantar) (Bulak et al. 2023 , Kellner and Sayão 2023 , Weinschutz et al. 2023 , Souza et al. 2023 ).
1.9. Other studies on prehistoric avifauna
Inferences about the Brazilian prehistoric avifauna were also made from other sources than physical remains. Yamashita ( 1997 ) hypothesized that Anodorhynchus macaws were followers of giant herbivorous mammals during the Pleistocene—dubbed as anachronistic survivors by Agnolín ( 2016b ), consuming the palm nuts that were stripped off the pulp by the herbivores, and this conservative and primitive behavior is now present again in their association with the domestic cattle introduced in their native range, which similarly affects the landscape. Among other factors, the Anodorhynchus material from the Lagoa Santa region, with its well-documented extinct mammalian megafauna, was used to support the hypothesis.
Owl pellets are common in quaternary sites, such as those mentioned by Lund ( 1839 , 1841b , 1841c , 1846 ) from Pleistocene–Holocene sites in the Lagoa Santa region in Minas Gerais, and the holocenic archeological sites Garivaldino, Pilger, and Sangão in Rio Grande do Sul (Fernández et al. 2019 ), likely produced by Tyto furcata . Furthermore, a holocenic mammalian bone assemblage possibly accumulated by vultures was reported by Ballejo et al. ( 2022 ) from Gruta do Presépio in Rio do Oeste, Santa Catarina. These pellets were found together with faunal remains discarded by humans that lived in the rock shelter from the early to the late Holocene and constitute the first fossil record of a bone assemblage accumulated by these birds.
Birds are a common motif in rock paintings from various traditions in Brazil. Naturally, most identifications are tentative, but in sites where multidisciplinary studies have taken place some paintings could be assigned to family or genus level. Rhea-like figures are abundant in Northeast rock shelters, such as those from the São Raimundo Nonato region (e.g., Guidon 1991 , Pessis 2003 , Faure et al. 2010b ), which also harbors figures interpreted as ciconiids, tinamids, falconids, and possibly anhimids (Schmitz 1990 ). Rheas, psittacids, and toucans were recorded in the Seridó region in Rio Grande do Norte (Schmitz 1990 ), and a flamingo-like figure was found in São Rafael (Souza and Medeiros 1982 ). Rheas and birds of prey are present in the Santa Elina rock shelter in Jangada, Mato Grosso (Vilhena Vialou and Vialou 1989 ), and rhea and psittacid-like figures were found in Serranópolis, Goiás (Dias 2004b , Schmitz 1990 ), among numerous others, often generic representations throughout the country (e.g., Silva et al. 2007 ). Besides rock paintings, other artistic manifestations add to this category, such as the zoomorphic figures made of stone and bone found in sambaquis, which include representations interpreted as kingfisher, penguin, albatross, owl, and king vulture (Gaspar 2004 ). A thorough, multidisciplinary review of these records will certainly contribute to the understanding of the biology, biogeography, and relationship with humans of late Pleistocene–early Holocene avifaunas of Brazil.
1.10. Osteological studies
Osteological studies are a fundamental source of information for avian comparative and functional anatomy, biomechanics, systematics, phylogeny, paleontology, and archeology (Sick 1997 , Olson 2003 , Höfling 2004 ). Unfortunately, the field is still developing at a slow pace in Brazil, despite showing indications of growth since the beginning of the 2000s (SAPE 2002 ). The lack of comparative collections is a determining factor (Alvarenga 1992 ), and so is the lack of avian anatomists, who work mainly with living taxa (Höfling 2004 ). Today, the country’s five largest bird skeleton collections are held, beginning from the largest, in MHNT, PUC Minas, MZUSP, MN, and MPEG. A significant part of the work developed in this field results from undergraduate and graduate programs (Borges 2008 , Mallet-Rodrigues 2016 ), especially at the Universidade de São Paulo (in São Paulo) and Universidade Estadual Paulista (in Botucatu).
The following groups are included among articles and theses (that were not published as articles) dealing with non-paleontological and non-veterinary osteology (excluding conference abstracts), with examples of studies: Tinamiformes (Silveira and Höfling 2007 ), Anseriformes (Previatto 2012 ), Galliformes (Silveira 2003 , Marceliano et al. 2007c ), Columbiformes (Andrela and Donatelli 1995 , Marceliano et al. 2007b ), Cuculiformes (Posso and Donatelli 2001 , 2005 , 2006 , 2007 , 2010 ), Nyctibiiformes (Mahecha and Oliveira 1998 , Höfling and Alvarenga 2001 , Costa and Donatelli 2009 , Costa 2014 , Costa et al. 2017 , 2021 ), Caprimulgiformes (Höfling and Alvarenga 2001 , Costa 2014 ), Apodiformes (Höfling and Alvarenga 2001 ), Opisthocomiformes (Marceliano 1996 ), Gruiformes (Marceliano et al. 1997 , Alves 2012b ), Eurypygiformes (Marceliano et al. 2007a ), Procellariiformes (Dénes and Silveira 2007 ), Ciconiiformes (Guzzi 2007 , Guzzi et al. 2014a , Santos et al. 2018 , Oliveira et al. 2019 ), Pelecaniformes (Ferreira and Donatelli 2005 , Ferreira 2007 , Silva 2011 ), Cathartiformes (Brito 2008 ), Accipitriformes (Migotto 2008 , 2013 ), Strigiformes (Mahecha and Oliveira 1998 , Höfling and Alvarenga 2001 , Salomão 2015 ), Coliiformes (Höfling and Alvarenga 2001 ), Trogoniformes (Höfling and Alvarenga 2001 ), Coraciiformes (Calonge-Méndez 1998 , Flausino et al. 1999 , Höfling and Alvarenga 2001 , Pascotto and Donatelli 2003 , Pascotto et al. 2006a , 2006b , Calonge-Méndez and Höfling 2007 ), Galbuliformes (Donatelli 1992 , Alvarenga et al. 2002 , Ladeira and Höfling 2007 , Posso et al. 2020 ), Piciformes (Höfling and Gasc 1984a , 1984b , Höfling 1991 , 1995 , Donatelli 1996 , 2012a , 2012b , Höfling and Alvarenga 2001 , Novaes 2013 , Araújo 2014 , Caldas et al. 2019 ), Falconiformes (Silva et al. 2012a , Guzzi et al. 2014b , 2015a , 2015b ), Psittaciformes (Porto 2004 , Gaban-Lima 2007 , Ferraroni 2015 ), and Passeriformes (Donatelli 1997 , Höfling and Alvarenga 2001 , Almeida 2003 , Calonge-Méndez 2004 , Donatelli and Marceliano 2007 , Cid 2011 , Araújo 2012 , Previatto and Posso 2015 , Guzzi et al. 2016 ). Additionally, Aires ( 2019 ) and Aires et al. ( 2021 ) worked with the osteology of both fossil and living birds.
2. CATALOG OF RECORDS
This section lists the records of fossil and subfossil material attributed to birds (as well as those of dubious affinity) from the literature and some that were unpublished.
2.1. Composition
The records are grouped here according to the political geographical unit established as the Brazilian territory, thus being an artificial composition. Chronologically, they range from the Early Cretaceous to recent, pre-colonial archeological material. They include bones, feathers, mummified specimens, eggs, coprolites, and tracks found in cave, tank, lake, fluvial, and coastal deposits, besides archeological contexts such as sambaquis and cerritos. Fossil feathers of uncertain determination, which may represent non-avian dinosaurs, were also included.
This study is not intended to be a complete catalog of all material unearthed in Brazil, since this is an impossible task, but rather the closest approximation allowed to this objective. Great efforts were made to cover as much paleontological and archeological literature as possible. However, since the archeological material is more sparsely mentioned in the literature, it is more likely that some records have not been included here. Taxa determined to the species level are most likely to be included, and, therefore, the coverage of this type of occurrence is biased, in addition to the existing biases regarding this kind of material inside archeological methods.
Considering the occurrence of certain groups in North America and southern South America, such as the Presbyornithidae (Mourer-Chauviré 1999 , Agnolín 2016b ), it is very likely that they also inhabited Brazil, although no fossils have been found so far. Alvarenga ( 1993b: 21), emphasizing the enormous wingspan (about 6 m) of Argentavis magnificens (Miocene of Argentina), commented that “it is reasonable to accept that this giant also flew over Brazilian territory”. Odontopterygiform birds, apart from a dubious record (see below), likewise inhabited the territory, considering they were recorded throughout the globe, with South American records in Chile, Peru, and Venezuela (e.g., Chávez et al. 2007 , Mayr and Rubilar-Rogers 2010 , Solórzano and Rincón 2015 ).
Despite some material being solely attributed to indeterminate birds, historical and noteworthy records will be discussed in proper accounts.
Fossil and subfossil pellets, like those accumulated by owls in several rock shelter sites, are not included in this review.
2.2. Notes on descriptions
Each record account is structured as follows: the general number in this study, the designation used in this study, the temporal (geochronologic unit) and spatial (Brazilian state) sources, the type locality (as in the original) and etymology for extinct described species, a chronological synonym list, and the descriptive text.
A “†” preceding a determination indicates the taxon is confirmed to be extinct. A “?” after the determination of a taxon indicates the attribution of such a name is uncertain. A “cf.” before a genus or species indicates the material is comparable, but not securely determined, to such taxon. An “aff.” before a species indicates the material is closely related to, but not identical, to such taxon. A “(spp.)” following a determination indicates that multiple indeterminate species-level taxa are included in that record. Different indeterminate records with the same designation (e.g., “Passeriformes indet.”) are numbered and ordered first geochronologically and then by their first mention in the literature.
The items in the synonym lists include the name the record was referred to and the author, year, and page (that describes the material or name it) of the publication (including theses and conference abstracts), besides figures (abbreviated as “fig.” or “figs”) and plates (abbreviated as “pl.” or “pls.”) when available. Tables are not included. Generally, the most exclusive term in a publication is used; vague terms such as “bones” or “feathers” are avoided when known to refer to several specimens previously mentioned with more exclusive names. The criteria for inclusion of names in the synonym lists are based on the most important publications within ornithology and paleontology, publications of historical importance, and/or publications that differ from the current dominant designation for such material. A new genus or species name described from Brazilian material is indicated by “[new genus and species]”, “[new genus]”, or “[new species]”. A name followed by “[in part]” indicates that such designation refers to only a part of the total material included in that record. A name followed by “[?]” indicates that such designation is tentatively attributed to that record. A designation followed by another between brackets indicates two different names used for the same material in the same study.
The descriptive texts gather information on the discovery process (e.g., locality, age, discoverer, descriptor), material content (e.g., known elements, holotypes and paratypes, state of preservation, housing institution, institutional identification), implications for diversity, ecology, evolution, classification, temporal and spatial distribution, figures, and other observations. Since Brazil’s bird fossil record and its history are fluid, and at the risk of being repetitive, we refrained from following a rigid scheme of standardization in these texts, as it would be sectioning them into topics. In-depth information about anatomical descriptions and measurements, taphonomic processes, and cultural aspects regarding zooarcheological remains should be consulted in the cited bibliography and references therein.
The osteological terminology adopted here follows Baumel and Witmer ( 1993 ), with terms adapted to English. Feather terminology follows the original descriptions.
Museum collection codes are kept up to date as much as possible. For example, old DGM codes have been replaced by newer MCT ones (e.g., DGM 1421-R is now MCT.R.1421).
2.3. Notes on nomenclature and taxonomy
Non-osteological records are arranged primarily geochronologically and, if available, taxonomically. Osteological records are arranged primarily taxonomically. Sub-records are arranged following the date of the first mention in the literature. The taxonomic nomenclature and systematic order adopted here follows Pacheco et al. ( 2021 ) for the Brazilian taxa and The Cornell Laboratory of Ornithology’s “Birds of the World” (Billerman et al. 2022 ) for taxa from elsewhere. Extinct taxa nomenclature follows specific authors, as seen in their respective texts. Extinct taxa are included chronologically after the living taxa’s systematic order.
The revised name attributions to the fossil materials follow largely on the lists of Lambrecht ( 1933 ), Brodkorb ( 1963 , 1964 , 1967 , 1971 , 1978 ), Mones ( 1986 ), and Cuello ( 1988 ), in addition to this study’s observations. However, these authors simplify most determinations given with uncertainties (such as those by Winge 1887 ), and some indeterminate material is not mentioned at all. In some cases, even records attributed to specific names (as some materials described by Winge) are lacking in Mones. These approaches are not adopted here. Lumping different records within the same uncertain determination (“indet.”, “cf.”) is not adopted here.
We also refrained from keeping records at the subspecies level, although some recent records could naturally be associated with the known subspecies in their origin area. This should be avoided since subspecies-defining methods generally do not include osteological characters, and much of the published material is only tentatively determined.
2.4. Records
The records herein are divided into four groups regarding the nature of their preservation: footprints, coprolites, feathers, eggs, and skeletal remains (including rarer mummifications). Refuted records are also commented on.
2.4.1. Refuted records
There are records in the literature that have been initially associated with birds, but further examination proved them to belong to other groups. Klein (1975 apud Ruschi 1979 ), Klein and Ferreira ( 1979 ), and Klein ( 1993 ) mentioned possible avian remains from the Itapecuru Formation (Early Cretaceous) of Maranhão. Through verbal communication with Diógenes de Almeida Campos, Martins Neto and Kellner ( 1988 ) stated these small-sized remains were being studied and did not belong to birds. Nevertheless, later mentions in the literature persisted (e.g., Santos and Carvalho 2004 ). Fernandes ( 2005 ) mentioned an isolated tetrapod footprint possibly belonging to Aves from the paleodesert of the Botucatu Formation (Late Jurassic–Early Cretaceous) in Araraquara, São Paulo, but this determination is now discarded (M.A. Fernandes, personal communication). Lopes et al. ( 2011 ) reported purported bird tracks from the Late Cretaceous of Serra do Tucano Formation in the Tacutu Basin. This material, however, was later attributed to Petroxestes ichnogenus of bivalve borings (Ioris 2015 ).
2.4.2. Footprints
So far, only one reliable record of avian footprint has been published for the country.
1. † Gruipeda isp.
Lima et al. ( 2023 ) reported an avian footprint from the shales of the Tremembé Formation. It is the first reliable record of this kind for the country. The fossil (UERJ IC-171) was collected in January 2023 during fieldwork carried out by the team of the Laboratório de Paleontologia of UERJ in an outcrop located at the Sociedade Extrativa Santa Fé in Tremembé. The footprint is 21 mm long and 18 mm wide, occurs in a level of grayish shale, and corresponds to a negative hyporelief with passive infilling of gray color that is lighter than the matrix color. Two elliptical coprolites of unspecified origin are present in the same sample. The footprint preserves features assigned to the four toes of the right foot, being characterized as mesaxonic and plantigrade. The authors tentatively assigned it to Gruipeda isp., and it was likely produced by members of Opisthocomidae, Rallidae, or Quercymegapodiidae.
2.4.3. Coprolites
Prehistoric avian coprolites have been only reported from the Tremembé Formation in São Paulo. Recent examples are known in the post-colonial period, such as those from Toca da Baixa dos Caboclos in Piauí (Sianto 2009 ).
2. †Coprolites 1 (MN 5619-I and MN 5620-I)
Late Oligocene/Early Miocene – SP
“Os nódulos coprolíticos teriam sido, pelo resultado descrito, produzidos por uma ave de porte médio de hábito alimentar múltiplo” – Castro et al. 1988a: 109.
“Coprólitos de aves” – Castro et al. 1988b: 2358–2370, figs 3–6.
“coprólitos de aves” – Carvalho and Fernandes 1989: 420–421, figs 3–4.
Neornithes [in part] – Carmo et al. 2024: 5.
Castro et al. ( 1988a , 1988b ) reported coprolites associated with birds from the shales (predominantly) and clays of the Tremembé Formation in the Fazenda Santa Fé. The coprolitic nodules are beige, allowing a good distinction from the matrix, with ellipsoidal (more frequent), circular, semicircular, or reniform shapes, and dimensions ranging from 6 to 50 mm along the major axis and commonly 5 to 20 mm along the minor axis. The material is deposited in the MN fossil invertebrate collection under numbers MN 5619-I (samples C-1 to C-6) and MN 5620-I (illustrated in figures 2–6 of Castro et al. 1988b and figures 3–4 of Carvalho and Fernandes 1989 ).
By analyzing its content and the chemical elements that make up the coprolitic nodules, it was possible to infer the feeding habits and the conditions in which its generators lived. Some of the nodules contain undigested ossicles and scales of characid fishes (Carvalho and Fernandes 1989 ), and elements that form a set of essential micronutrients to the development of plants were found, which corroborated the variation in these birds’ feeding habits. Carvalho and Fernandes ( 1989 ) postulated they might have originated from rallids, anatids, phalacrocoracids, or ardeids, who inhabited the border of the paleolake surrounded with riparian forest. The indication that these birds were not exclusive fish consumers agrees with the unfavorable environmental conditions to this type of diet, where periodic fish deaths were caused by the lake bottom’s anoxic characteristics, which limited their availability. The different diets of fossil birds known from the Tremembé Formation (from generalists to exclusive fish and carrion feeders) demonstrate the relationship between the chemical composition of the coprolites and the habits of the organisms that generated them.
The type of deposition of the coprolites suggests they were projected vertically in shallow water. Its occurrence corroborates the tendency of the drying up of the Tremembé paleolake, also evidenced by the accumulation of fish, traces of invertebrates, and vegetation in the swampy environment, until its complete drying.
3. †Coprolites 2
Late Oligocene/Early Miocene – SP
“bird coprolite” – Souto 2012: 112, 5C.
Souto ( 2012 ) depicted a coprolite associated with a bird from the Taubaté Basin, different from those described by Castro et al. ( 1988a , 1988b ) and Carvalho and Fernandes ( 1989 ).
4. †Coprolites 3 (UERJ IC-58)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Plant fragments (possibly Poaceae) were found in a sample extracted from it.
5. †Coprolites 4 (UERJ IC-59)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
6. †Coprolites 5 (UERJ IC-60)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
7. †Coprolites 6 (UERJ IC-61)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
8. †Coprolites 7 (UERJ IC-62)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
9. †Coprolites 8 (UERJ IC-63)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. One Trichocephalida egg and fish bone fragments were found in a sample extracted from it.
10. †Coprolites 9 (UERJ IC-65)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Plant fragments were found in a sample extracted from it.
11. †Coprolites 10 (UERJ IC-66)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
12. †Coprolites 11 (UERJ IC-67)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
13. †Coprolites 12 (UERJ IC-68)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
14. †Coprolites 13 (UERJ IC-69)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Two eggs, one of Ascaridina (morphotype I) and one of an indeterminate nematode, as well as fragments of fish bones and arthropods were found in a sample extracted from it.
15. †Coprolites 14 (UERJ IC-70)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
16. †Coprolites 15 (UERJ IC-71)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Two Spirurina eggs and fish bone fragments were found in a sample extracted from it.
17. †Coprolites 16 (UERJ IC-72)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
18. †Coprolites 17 (UERJ IC-73)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Fish bone fragments were found in a sample extracted from it.
19. †Coprolites 18 (UERJ IC-74)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Arthropod fragments were found in a sample extracted from it.
20. †Coprolites 19 (UERJ IC-75)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
21. †Coprolites 20 (UERJ IC-76)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé.
22. †Coprolites 21 (UERJ IC-77)
A coprolite reported by Carmo et al. ( 2023 ) from the Tremembé Formation shales of the Fazenda Santa Fé. Ascaridina eggs (3 of morphotype II and 12 of morphotype III) and fish bone fragments were found in a sample extracted from it.
2.4.4. Feathers
Fossil feathers are usually of little taxonomic importance but can demonstrate the presence of birds in certain deposits (Martill and Filgueira 1994 ), even though, regarding Mesozoic sediments, they could belong to other feathered dinosaur clades. Due to their fragility and the requirement of special conditions that allow their fossilization (Kellner et al. 1994 ), the rarity of such structures in the fossil record makes it worthwhile to comment on these records here. Specimens preserving coloration patterns and possible remains of parasites are of particular importance. Furthermore, recent avian feathers were described in the archeological literature, such as a psittacid feather from Toca do Sítio do Meio in Piauí (Melo 2007 ).
2.4.4.1. Mesozoic feathers
23. †Fossil feather 1 (GP/2T-136)
Early Cretaceous – CE
“Pena” – Martins Neto and Kellner 1988: 61–68, figs 1A, 1B, 2E.
“Neornithes ? ou Enantiornithes ?” – Viana and Mamede 1990: 486.
“isolated feather” – Chiappe 1991: 61.
“Undetermined feather” – Kellner et al. 1991: 376, fig. in pp. 377.
“feather, family incertae sedis ” – Martill 1993: 150.
“small flight feather” – Martill and Frey 1995: 121.
“flight feather” – Chiappe 1996: 535, 538–539, 3A.
“penas de diferentes formas e tamanho” [in part] – Kellner 1998: 657.
“Flight feather” – Kellner and Campos 2000: 523, 527, 19.
“markedly asymmetrical wing contour feather” – Martill and Davis 2001: 252.
“flight feather” – Kellner 2002: 396, 16.9.
“flight feathers” [sic] – Machado and Kellner 2007: 300.
“Asymmetrical remex” – Naish et al. 2007: 526, 18.1A.
cf. Aves [in part] – Bittencourt and Langer 2011: 28, 38.
“Penas isoladas – de contorno, das asas, da cauda e semi-plumas” [in part] – Campos 2011: 90.
“small flight feather” – Sayão et al. 2011: 198, 203.
“provável pena remex” – Leite and Hessel 2011: 2.
“One primary remex” – Prado et al. 2016a: 4.
Martins Neto and Kellner ( 1988 ) reported a feather (GP/2T-136) from the Crato Formation, which is the first record of this kind for the Mesozoic of South America and the first purported record of Mesozoic bird remain from Brazil. The relatively well-preserved fossil was provided to one of the authors by Pedro Luiz Novaes Ferreira, from São Paulo, and, although its exact provenance is unknown, it was likely collected around Santana do Cariri, Ceará. According to Chiappe ( 1996 ) through communication with Kellner, it comes from an indeterminate area around Nova Olinda.
The feather shows limonitic/imprint preservation (Prado et al. 2016a ). Unfortunately, the slab was fractured into several pieces, with one of the fractures dividing the fossil into two parts. The material reached the authors already glued, and, due to its fragility, it could not undergo further preparation. It is asymmetric with identifiable calamus (about 5 mm long), rachis, and barb impressions (about 19 barbs per centimeter of rachis), with a total length of about 64 mm, and a maximum width of about 8 mm. No barbule impressions are preserved, but due to the disposition of the barbs, the authors concluded they existed and were not preserved.
The fossil was attributed to a bird that was possibly well-adapted to flight due to the rachis curvature and vane asymmetry, probably a primary remex of a neornithean or an enantiornithean. Chiappe ( 1991 ) suggested it could belong to an enantiornithean, although there is no direct supporting evidence. Later, Barrowclough (in Kellner et al. 1991 ) suggested it may represent either one of the trailing primaries or a secondary remex, and if it was indeed capable of flight, the bird had approximately the size of a brushfinch and perhaps weighed between 35 and 40 g. A horizontally flipped, color picture of the specimen was included in the same publication. Sayão et al. ( 2011 ) commented that the feather could have belonged to a flightless form due to its similarity to that of flightless birds, as in the absence of barbules, although, as noted by Martins Neto and Kellner, this could be a preservation artifact.
24. †Fossil feather 2 (LEIUG 114369)
Early Cretaceous – CE
“Feather” – Martill and Filgueira 1994: 483–487, 2.
“demiplume” [sic] – Martill and Frey 1995: 121.
“semiplume” – Chiappe 1996: 539.
“Semiplume” – Kellner and Campos 2000: 523, 528.
“small semiplume” – Kellner 2002: 396–397.
“semiplume” – Machado and Kellner 2007: 300.
“semiplume” – Naish et al. 2007: 525.
“Several types of feathers” [in part] – Bittencourt and Langer 2011: 28.
“Penas isoladas – de contorno, das asas, da cauda e semi-plumas” [in part] – Campos 2011: 90.
“two small semiplumes” [in part] – Sayão et al. 2011: 198, 202.
“semipluma” – Leite and Hessel 2011: 2.
“Semiplume” – Prado et al. 2016a: 4.
Martill and Filgueira ( 1994 ) reported a feather (LEIUG 114369) from the Crato Formation of Mina de Antone Phillipe near the Tatajuba Reservoir, collected by one of the quarry workers.
The fossil is preserved as a carbonized trace (Prado et al. 2016a ) and was damaged since or during the collection. It is a semiplume associated with a bird, lacking the proximal part of the calamus. The preserved impression is 21 mm long along the slightly curved rachis and has a maximum width of 18 mm, partly due to the spreading of the loose and plumaceous barbs. The longest barbs range from 8 to 10 mm in length. Some are separated from adjacent barbs, with some folded back toward the proximal end of the rachis. Barbules, both proximal and distal, are identifiable in some barbs, being up to 0.04 mm long with up to sixteen barbules per millimeter in the parts of the barbs where counting them is easy. The feather is overall dark gray distally to light gray proximally.
It is similar to the feathers found in the posterior part of the body of modern passeriforms [18] and has a size comparable to those of birds between 150 and 300 mm long. The authors noted a remarkable similarity in size, morphology, and preservation with a fossil feather depicted by Talent et al. ( 1966 ) from the Early Cretaceous of Koonwarra, Victoria, Australia.
25. †Fossil feather 3 (MCT 1493-R)
Early Cretaceous – CE
“Fossil down feather” – Kellner et al. 1994: 489–492, 2.
“down feather” – Chiappe 1996: 538–539.
“penas de diferentes formas e tamanho” [in part] – Kellner 1998: 657.
“Down feather” – Kellner and Campos 2000: 523, 528, 17.
“very small and fluffy down feather” – Kellner 2002: 397, 16.10.
“Pena fóssil” – Kellner 2006: 84, 46.
“down feather” – Machado and Kellner 2007: 300.
“isolated down feather” – Naish et al. 2007: 525.
“Several types of feathers” [in part] – Bittencourt and Langer 2011: 28.
“Penas isoladas – de contorno, das asas, da cauda e semi-plumas” [?; in part] – Campos 2011: 90.
“down feather” – Sayão et al. 2011: 198.
“pluma” – Leite and Hessel 2011: 2.
“Down feather” – Prado et al. 2016a: 4.
A down feather (MCT 1493-R) [19] associated with a bird was described by Kellner et al. ( 1994 ) from the Crato Formation, without attribution to a specific locality. According to Chiappe ( 1996 ) through communication with Kellner, it is from an indeterminate area around Nova Olinda.
It is a comparatively small feather, preserved as a carbonized trace (Prado et al. 2016a ), with a length of about 7.5 mm from the faintly visible calamus to the largest barb, and is of a dark-brown color in a grayish slab. The very slender rachis is about 3.4 mm long. Some barbs are almost as thick as the rachis. They show individual sizes that decrease towards the shaft’s distal end, with the last and smallest barb on the shaft’s terminal portion about 2.9 mm long, while the largest barb near the shaft’s base is about 5.3 mm long. There are about three to four barbs per millimeter of the rachis. Barbules are longer on the barbs’ proximal parts, increasing the down density of the feather towards the rachis. On the upper left portion of the matrix above the feather there is a long barb-like structure, at least 25 mm in length, straight distally and curved proximally. It was possibly originally connected to the largest basal barb or not part of the feather at all, as no barbules were observed in the structure.
26. †Fossil feather 4 (LEIUG 115562 and SMNK 1247 PAL)
Early Cretaceous – CE
“small ?breast feather” – Martill and Frey 1995: 120–121, 1.
“contour, body feather” – Chiappe 1996: 539.
“Contour feathers” [in part] – Kellner and Campos 2000: 523, 528.
“incomplete contour feather” – Kellner 2002: 397.
“contour feathers showing part of the color pattern” [in part] – Machado and Kellner 2007: 300.
“colour-banded feather” [in part] – Vinther et al. 2008: 522–525, 1A–C.
cf. Aves [in part] – Bittencourt and Langer 2011: 28, 38.
“Penas isoladas – de contorno, das asas, da cauda e semi-plumas” [?; in part] – Campos 2011: 90.
“contour feathers” [in part] – Sayão et al. 2011: 198, 202.
“pena de contorno” – Leite and Hessel 2011: 2.
“Contour feather” – Prado et al. 2016a: 4.
“striped fossil feather” [in part] – Smithwick and Vinther 2020: 190–191, 11.5.
“feather” [in part] – Roy et al. 2020: 254.
Martill and Frey ( 1995 ) reported a feather with a preserved color pattern from Mina de Antone Phillipe near Tatajuba Reservoir. The part (LEIUG 115562) and the counterpart (SMNK 1247 PAL) were collected and deposited in different institutions.
It is a small body feather, possibly from the breast, 7 mm long with a maximum width of 4 mm, from a bird probably slightly larger than a living fringillid. The specimen was split in half, with the color pattern preserved on both part and counterpart. It comprises a series of transverse bands of alternating dark and light appearance with relatively sharp limits between the colored bands. Three dark bands exist perpendicularly to the rachis, the most proximal one beginning at the quill’s distal portion. The proximal portion of the feather is partially concealed by the matrix, concealing the entirety of the quill’s length in LEIUG 115562, which also has bands that appear overall darker. A small distal extension of color exists along the rachis, but it does not reach the succeeding dark color band. A dark band includes the feather’s tip, where there is also a proximal extension of the band along the rachis that almost unites with the distal extension of the central dark band. The authors commented that this barred feather pattern probably gave the bird a speckled appearance, such as seen in the breast of some turdids and fringillids. According to Kellner ( 2002 ), it most likely represents only the apex of the pennaceous portion of a body feather.
A study conducted by Vinther et al. ( 2008 ) demonstrated that the dark bands of LEIUG 115562 are preserved as elongate, oblate carbonaceous bodies 1–2 μm long, aligned along with the barbs and barbules, whereas the light bands retained only relief traces in the matrix. These oblate bodies were interpreted as eumelanosomes, as seen in the structure of black feathers in the living icterid Agelaius phoeniceus . Their observations indicated the specimen was originally banded with a black-and-white eumelanin pattern.
27. †Fossil feather 5 (LEIUG 115563 and SMNK 1251 PAL)
Martill and Frey ( 1995 ) reported a second feather from the same locality as the specimen above. It is about 15 mm long and 1.5 to 2.0 mm wide, as all the barbs are pressed together. The authors selected this second feather for examination through electron microscopy to avoid damaging specimens (including insects) with a preserved color pattern, as coating with gold or carbon would mask the preserved pattern. This second feather is preserved similarly to the color-patterned feather but lacks this pattern, possibly because it had an original uniform color. The specimen was split in the middle, with half the feather remaining on each counterpart slab. One half remained preserved (SMNK 1251 PAL) while the other (LEIUG 115563) was trimmed with a diamond saw so it could be mounted on a flat-topped aluminum stub and later sputter coated with gold and examined in a scanning electron microscope. At high magnifications, it showed that the feather’s organic material is composed of roughly aligned elongate, rod-shaped bodies 0.5–2.2 μm long with a diameter of 0.3–0.5 μm that the authors considered to be autolithified bacteria, resembling that from the Eocene oil shales of Messel, Germany. The feather per se was not depicted, but bacteria micrographs were included in the study. However, as noted above, Vinther et al. ( 2008 ) interpreted similar structures in the banded feather described from the same locality as melanosomes.
28. †Fossil feather 6 (NSM PV 20059)
Early Cretaceous – CE
“fossil feather” – Martill and Davis 1998: 528–529, 1.
“feather” – Martill and Davis 2001: 241–259, figs 1–5.
“isolated feather with a symmetrical vane” – Naish et al. 2007: 526, 528, 18.1B–F.
“Penas isoladas – de contorno, das asas, da cauda e semi-plumas” [in part] – Campos 2011: 90.
“possible rectrices” [in part] – Sayão et al. 2011: 202.
“pena de contorno” – Leite and Hessel 2011: 2.
“One symmetrical feather” – Prado et al. 2016a: 4.
“rêmige” – Prado 2017: 75.
Martill and Davis ( 1998 ) described a feather with possible ectoparasite eggs (NSM PV 20059) from the Crato Formation and later elaborated on it further (Martill and Davis 2001 ). The National Science Museum of Japan purchased the specimen from a fossil dealer in June 1996. Although the locality was given only as Crato Formation, the authors confirmed that the specimen matrix is similar to that of weathered, laminated limestones of the Nova Olinda Member, highly likely from one of the small quarries between Nova Olinda and Santana do Cariri. Although seemingly preserved as a carbonized trace, the authors noted that the fossil is too weathered to establish its original preservation mode satisfactorily, but it was considered so by Prado et al. ( 2016a ).
The fossil comprises part and counterpart (this one less preserved), orange/brown in coloration on a buff-colored slab. The feather is 85 mm long and has a maximum width of 11 mm, while the calamus is 19 mm long and 1.5 mm at its widest part. The umbilical sulcus is possibly represented by a 33 mm long dark-colored band lying centrally within the scapus. The rachis (which the authors considered part of the scapus bordered by the barbs) is 66 mm long and slightly curved. In the apex, possibly two or three millimeters are missing. The rachis’ margins seem to diverge proximally at about 48 mm from the distal apex, possibly being the termination of the rachis part occupied by medullary tissue combined with the taphonomic artifact of the rachis collapse along the umbilical sulcus from a three-dimensional state to the fossil’s almost two-dimensional state. Although there is a real slight asymmetry, the main slab specimen shows a visually apparent asymmetry due to the barbs being damaged distally on the right vane’s margin. The maximum length of the barbs is 17 mm at a point about 35 mm from the proximal umbilicus at the proximal tip of the calamus. Although not well-preserved, the barbules are visible on many of the barbs. They are single, fine distolateral projections, about 0.25 mm long, in the barbs’ proximal and distal margins, where they appear to be densely distributed. A narrow space running between the barbs results from the apparent lack of interaction between barbules of adjacent barbs in most regions of the vane. The feather is almost complete. It seems to lack only very small portions of the distal tips of the most distal few barbs and shows some damage distally in some of the more proximal barbs.
Adhered to the barbs, barbules, and calamus are numerous, spherical to sub-spherical egg-like structures, which the authors ruled out as artifacts of preservation or a pollen affinity in favor of the identity as true eggs. They are reddish-brown and slightly shiny, present over most of the feather’s surface, with a minimum number of 242 units on the most well-preserved slab. They have a diameter of 68–75 μm and a wall of 5–7 μm thick in split units that allow measurement, and occur mainly in isolation, although a few irregular clusters of between 5 and 15 units are also present. The structures were hollow (now filled with calcite), and some display a circular aperture on the surface facing away from the feather, which suggests they had hatched (Naish et al. 2007 ). The association between the feather and eggs is not unequivocal, and possibly they were from a microscavenger rather than a parasite. However, their number and close resemblance to the eggs of modern bird mites suggest they are at least eggs of mites (Acari) if not parasitic mites. Proctor ( 2003 ) suggested that if these structures are indeed eggs, they may be from ostracods, and argued that modern feather mite eggs are larger (150–400 μm long), sausage-shaped, and have longitudinal seams rather than circular apertures, and ostracods readily deposit their small, round eggs on submerged detritus. While there are no records of ostracods in the Nova Olinda Member, they occur in abundance in the strata immediately beneath (Naish et al. 2007 ). Prado ( 2017 ) reinforced this hypothesis by mentioning the significant difference in the distribution pattern of modern pennaceous feather parasites.
The feather is possibly precluded from being a flight feather due to its near symmetry and is similar in this aspect to the rectrices of Archaeopteryx lithographica . If a primary wing feather, it would be similar in asymmetry range to recent flightless birds. The barbs are straight for most of their length, but hooked barbules are absent, resembling those in the elongated body feathers of large ratites such as Rhea . The authors suggested that the specimen may have belonged to an animal with similar ecology, a ground-dwelling avian or feathered non-avian dinosaur but did not assign it to any group.
The feather’s isolated nature suggests it was possibly molted, being slightly damaged due to abrasion sustained in life (as in preening) or natural wear. The authors suggested the feather was self-plucked due to severe infestation based on the several supposed eggs on its surface being unhatched and many of the same deposit fossils appearing to have been blown into the Crato lagoon.
29. †Fossil feather 7 (MCT 1509-R)
Early Cretaceous – CE
“Pena fóssil” – Kellner et al. 1999: 53, 41.
“Contour feather” – Kellner and Campos 2000: 523, 528, 18.
“complete contour feather” – Kellner 2002: 397, 16.11.
“contour feathers showing part of the color pattern” [in part] – Machado and Kellner 2007: 300.
cf. Aves [in part] – Bittencourt and Langer 2011: 28, 38.
“Penas isoladas – de contorno, das asas, da cauda e semi-plumas” [?; in part] – Campos 2011: 90.
“contour feathers” [in part] – Sayão et al. 2011: 198, 202.
Kellner ( 2002 ) described a complete contour feather (MCT 1509-R), previously depicted by Kellner et al. ( 1999 ) and Kellner and Campos ( 2000 ), from a quarry of the Crato Formation near Nova Olinda. It is possibly a body feather, with a length of almost 22 mm. The rachis is curved, the vanes are symmetrical, and the proximal portion of the calamus was not preserved. Five pairs of alternating dark and light bands, essentially perpendicular to the rachis, compose the feather’s color pattern and are more visible toward the apex of the pennaceous part and fading towards the basal portion, which lacks any banding.
30. †Fossil feather 8 (Chiappe 2007 )
Early Cretaceous – CE
“Some fossil feathers” [in part] – Chiappe 2007: 78, fig. in pp. 78.
Chiappe ( 2007 ) depicted a 5 cm-long feather with a preserved banded color pattern and only stated that it comes from the Early Cretaceous of Brazil. It is similar to the specimens depicted by Kellner ( 2002 , 16.11) and Naish et al. ( 2007 , pl. 25E).
31. †Fossil feather 9 (Naish et al. 2007 1)
Early Cretaceous – CE
“isolated down feathers” [in part] – Naish et al. 2007: 528, pl. 25A.
Naish et al. ( 2007 ) depicted a down feather about 18 mm long from the Crato Formation. It is of dark color, and its distal half barbs are preserved clumped together.
32. †Fossil feather 10 (Naish et al. 2007 2)
Early Cretaceous – CE
“isolated down feathers” [in part] – Naish et al. 2007: 528, pl. 25B.
Naish et al. ( 2007 ) depicted a down feather about 18 mm long and 14 mm wide (due to the spreading of the barbs) from the Crato Formation. A darker tone is present in the distal barbs and the middle barbs’ distal parts.
33. †Fossil feather 11 (Naish et al. 2007 3)
Early Cretaceous – CE
“isolated down feathers” [in part] – Naish et al. 2007: 528, pl. 25C.
Naish et al. ( 2007 ) depicted a down feather about 17 mm long and 12 mm wide (due to the spreading of the barbs) from the Crato Formation. It has an overall dark coloration.
34. †Fossil feather 12 (Naish et al. 2007 4)
Early Cretaceous – CE
“small down feather with pattern of transverse bands” – Naish et al. 2007: 528, pl. 25E.
Naish et al. ( 2007 ) depicted a small down feather about 18 mm long from the Crato Formation. It preserves a pattern of transverse bands, similar to specimens depicted by Kellner ( 2002 , 16.11) and Chiappe ( 2007 , fig. in pp. 78).
35. †Fossil feather 13 (Naish et al. 2007 5)
Early Cretaceous – CE
“single elongate down feather” – Naish et al. 2007: 528, pl. 25F.
Naish et al. ( 2007 ) depicted an elongate down feather about 19 mm long from the Crato Formation.
36. †Fossil feather 14 (Naish et al. 2007 6)
Early Cretaceous – CE
“feather with pattern of fine, diagonally arranged banding” – Naish et al. 2007: 528, pl. 25G.
Naish et al. ( 2007 ) depicted a feather preserving a fine, diagonally arranged banding pattern, with a length of about 18 mm.
37. †Fossil feather 15 (Naish et al. 2007 7)
Naish et al. ( 2007 ) depicted a well-preserved, elongate, symmetrical feather with a pattern of offset bands (about 10 dark bands are visible in the right vane, fading towards the calamus), with a length of about 18 mm. The specimen was later depicted by Herzog et al. ( 2008 ), who commented that no information was available on the collection where the fossil was deposited (probably in Europe).
38. †Fossil feather 16 (CAV-0001-V)
Early Cretaceous – CE
“plúmula” – Sayão and Uejima 2009: 22.
“plúmula” – Sayão and Uejima 2010: 54.
“penas plumáceas” [in part] – Freitas et al. 2010: 32.
“penas plumáceas” [in part] – Freitas et al. 2011: 124.
“down feather” – Sayão et al. 2011: 199, 2.
“plumas” [in part] – Leite and Hessel 2011: 2.
“One plume” – Prado et al. 2016a: 4.
“CAV-001-V” – França et al. 2017: 19.
Two feathers collected during the field season of 2009 by biology student Laiz Karla of CAV at the Mina do Demar, on the road connecting Nova Olinda and Santana do Cariri, were briefly described by Sayão and Uejima ( 2009 , 2010 ) and depicted by Sayão et al. ( 2011 ). They are presumably preserved as carbonized traces (Prado et al. 2016a ). The first is a down feather (CAV 0001-V), which has a total length of 18.81 mm (excluding the calamus, which was not preserved) and a maximum width of 13.66 mm. The rachis is 5.07 mm long and is shorter than the barbs. The largest barb is 11.85 mm long, while the smallest is 5.4 mm long. The barbules are more abundant in the base of the barb and decrease in number and size distally. França et al. ( 2017 ) reported the rachis is 11 mm long and the largest barb 11.03 mm long.
39. †Fossil feather 17 (CAV-0002-V)
Early Cretaceous – CE
“semi-plúmula” – Sayão and Uejima 2009: 22.
“semi-plúmula” – Sayão and Uejima 2010: 54.
“penas plumáceas” [in part] – Freitas et al. 2010: 32.
“penas plumáceas” [in part] – Freitas et al. 2011: 124.
“semiplume” – Sayão et al. 2011: 199, 201, 3.
“plumas” [in part] – Leite and Hessel 2011: 2.
“One semiplume” – Prado et al. 2016a: 4.
“CAV-002-V” – França et al. 2017: 19.
The second feather from Mina do Demar described by Sayão and Uejima ( 2009 , 2010 ), and later depicted by Sayão et al. ( 2011 ), is a semiplume (CAV 0002-V). It is preserved in a dark brown color contrasting with the light-yellow matrix. A variation in the staining tone is present, darker in the base of the calamus and lighter closer to the rachis. A very light band pattern can be observed distally in the barbs, interposed between light and dark shades. The feather has a total preserved length of 8.55 mm. The calamus is 0.47 mm long and has been displaced from its original position, likely during fossilization. The very slender rachis is 4.76 mm long, longer than the barbs. The barbs decrease in size toward the shaft’s distal part. The largest barb is 4.71 mm long, while the smallest is 3.34 mm long. Both proximal and distal barbules are identifiable in some barbs. It was reported as the smallest semiplume described from the Crato Formation. França et al. ( 2017 ) reported the rachis is 7.49 mm long and the largest barb 4.91 mm long.
40. †Fossil feather 18 (Teixeira and Saraiva 2010 )
Teixeira and Saraiva ( 2010 , 2011 ), two researchers from URCA, reported a feather of a bird or a non-avian theropod among material collected in quarries of Nova Olinda.
41. †Fossil feather 19 (MPSC-PN 2221)
A third feather (MPSC-PN 2221) from Mina do Demar was described by Sayão et al. ( 2011 ). The feather possesses a long calamus and a rachis (also somewhat flattened) with a total length of about 75 mm. The loose barbs are long and filamentous and lack barbules. It is overall similar to down or ornamental feathers but differs from them in the rachis morphology. The authors also noted the similarity to a fossil feather preserved in amber from the Early Cretaceous of France described by Perrichot et al. ( 2008 ) and suggested a non-avian origin based on comparison with the developmental stages defined by Prum ( 1999 ). Conversely, Prado et al. ( 2016a ) mentioned it under “feathers assigned to a bird” preserved as carbonized traces in their Table 1 and mentioned the two other specimens first reported by Sayão and Uejima ( 2009 ) separately.
42. †Fossil feather 20 (UFC.0022V)
Leite and Hessel ( 2011 ) reported eight dark-colored feathers from Mina do Triunfo in Nova Olinda, which they tentatively attributed to contour feathers from the upper hindlimbs of non-avian dinosaurs. They noted similarities with the feathers from the Early Cretaceous of Liaoning, China described by Zhang et al. ( 2006 ). The slender rachis has almost the same width as the barbs, attached to the rachis at about 15°. The barbules are attached with a small angle and almost coalesce with the barbs forming a kind of sheath, especially in the distal barbs. One of these specimens is an isolated, apparently symmetrical feather (UFC.0022V) 22 mm long.
43. †Fossil feathers 21 (UFC.0023Va and UFC.0023Vb [MN 7754-V])
The other feathers reported by Leite and Hessel ( 2011 ) from Mina do Triunfo are a set of seven feathers in the same matrix block, preserved in part and counterpart (UFC.0023Va and UFC.0023Vb). They are almost symmetrical and range from 15 to 25 mm long. The feathers are united by their basal region and no traces of calami could be observed, but this area preserved a finely punctate epidermis impression of about 2 mm 2 .
Campos et al. ( 2019 ) analyzed the specimen through combined microscopy and spectroscopy techniques (they were aiming at minimal damage, only 1 mm³ was scrapped from the surface) and confirmed the fossils’ feather identity, distinguishing six (not seven) plumulaceous down feathers. The analysis also revealed preserved eumelanosomes with a mean length of 1.293±0.317 μm and a mean width of 0.298±0.032 μm in its composition, indicating the feathers were originally dark-colored. Barbules similar to that of extant plumulaceous feathers and long prong barbicels were also distinguishable. These conclusions should also be valid for the UFC.0022V specimen, as it is of the same aspect. Strangely, the set of feathers is referred to as MN 7754-V, and no further information was provided, nor Leite and Hessel were cited. It was also not specified if it was the part or counterpart (as such a feature is not mentioned), though the pictured unit from both studies matches well.
44. †Fossil feather 22 (GP/2E-7853)
Prado and Anelli ( 2013 ) briefly described two feathers from the Crato Formation, apprehended by the Polícia Federal, which were further mentioned by Prado and Anelli ( 2014b ), and described and referred to Coelurosauria by Prado et al. ( 2016a ).
The first is a down feather (GP/2E-7853) of orange coloration in a weathered beige slab, possibly preserved as limonite. A nearly complete Dastilbe sp. is present in the same matrix block, as well as ostracod remains (Prado 2017 ). The feather has a width of 12.36 mm and a length (excluding the calamus) of 16.14 mm. The calamus was not preserved and was supposedly very thin. The rachis is 0.49 mm wide and 9.43 mm long and has barbs of varying sizes, ranging from 4.85 mm to 8.65 mm. Barbules were only subtly preserved in some regions of the barbs.
Prado et al. tentatively assigned both this and GP/2E-7854 specimens to stage IIIb of Prum and Brush’s ( 2002 ) evolutionary model and to morphotype 4 of Xu and Guo ( 2009 ) and noted the possibility they were auricular feathers of an animal of a maximum size similar to that of a chicken. Due to the absence of melanosomes preserved in the fossil, Prado ( 2017 ) suggested the feather was either composed of other biochromes susceptible to degradation or unpigmented (white or cream-colored).
45. †Fossil feather 23 (GP/2E-7854)
The second feather described by Prado and Anelli ( 2013 , 2014b ) and Prado et al. ( 2016a ) is a down feather (GP/2E-7854) of varying brownish tones (due to different preservation in carbonaceous traces) in a weathered beige slab. It has a width of 12.76 mm and a length of 19 mm. The thin calamus, inferred from a slight-line structure consisting of an external molt, is 0.24 mm long. The rachis is 0.49 mm wide and 12.03 mm long, and the barbs range from 4.3 mm to 17.83 mm. The barbules are present as vestigial traces.
46. †Fossil feather 24 (GP/2E-8771)
In addition to the previous two apprehended specimens, Prado et al. ( 2016a ) described a semiplume (GP/2E-8771) of blackish color, possibly preserved as a carbonized trace in a grayish (likely unweathered) slab. It has a width of 15.63 mm and a length of 33.50 mm. The rachis is 0.03 mm wide and 29.35 mm long, and the barbs range from 4.12 mm to 16.45 mm. The barbules are clearly visible and vary in size, suggesting some degree of cohesion between the barbs, though no barbicels can be observed. A V-shaped structure reminiscent of afterfeather, larger than the vanes, is attached to the basal part, though it does not show afterfeather diagnostic features. Markings done with a scraper tool, especially in the region where the calamus was supposed to be found, were made by illegal dealers to make it more commercially attractive. Ostracod remains are present in the matrix (Prado 2017 ). The authors referred the specimen to Maniraptoriformes and assigned it to stage IIIa+b of Prum and Brush’s ( 2002 ) evolutionary model and morphotype 6 of Xu and Guo ( 2009 ). They also suggested it might have had protective and thermoregulatory functions and speculated on camouflage, communication, and sexual roles.
Feathers from the Crato Formation: GP/2E-7853 (A); GP/2E-7854 (B); GP/2E-9378 (C); GP/2E-9381 (D); GP/2E-8771 (E); GP/2E-9379 (F); GP/2E-9405 (G); GP/2E-9380 (H); GP/2E-9382 (I); GP/2E-9383 (J); GP/2E-9384 (K); GP/2E-9428 (L); GP/2E-9385 (M); GP/2E-9386 (N); GP/2E-9387 (O); GP/2E-9406 (P) (R.S. Nascimento). Scale bar: 5 mm.
Previously, Prado and Anelli ( 2015 ) performed SEM analysis in the specimen and the observed microbodies were interpreted as eumelanosomes, which suggested it had an iridescent pattern and the dinosaur fauna of that unit was well adapted to its ecological niche. Furthermore, Prado ( 2017 ) reported that the eumelanosomes’ morphology in this specimen suggests it was dark or slightly dark-colored, possibly grayish.
47. †Fossil feather 25 (UFRJ-DG 05 Av)
Early Cretaceous – CE
“UFRJ-DG 05 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It figures among the 45 specimens from Mina Pedra Branca in Nova Olinda collected between 2000 and 2016. It has a maximum width of 2.9 mm. In general, the contour feathers he described have pennaceous vanes, formed by parallel barbs with intertwined barbules. The calami are short and thin, difficult to distinguish in the matrix, and the rachises are preserved as a thin dark line between the vanes. The size of the plumulaceous areas varies.
48. †Fossil feather 26 (UFRJ-DG 08 Av)
Early Cretaceous – CE
“UFRJ-DG 08 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 20.9 mm, a maximum width of 14.9 mm, a rachis 14.76 mm long, and the longest barb 15.1 mm long.
49. †Fossil feather 27 (UFRJ-DG 09 Av)
Early Cretaceous – CE
“UFRJ-DG 09 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). It has a maximum length of 16.7 mm, a maximum width of 10.9 mm, the rachis is 11.3 mm long, and the longest barb is 11.4 mm long. It is curved, with loose barbs at the feather edge, similar to wing coverts of modern birds.
50. †Fossil feather 28 (UFRJ-DG 10 Av)
Early Cretaceous – CE
“UFRJ-DG 10 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 10.8 mm, a maximum width of 11.9 mm, the rachis is 3.2 mm long, and the longest barb is 7.9 mm long. In general, the plumes he described have slender rachis (not always easily recognizable), poorly preserved calami, and vanes formed by long and slender disarranged barbs with non-intertwined barbules that show a fuzzy texture. The longest barbs are longer than their respective rachises.
51. †Fossil feather 29 (UFRJ-DG 11 Av)
Early Cretaceous – CE
“UFRJ-DG 11 Av” – Metello 2017: 19–20, 10.
A contour feather described by Metello ( 2017 ). It has a maximum length of 14.4 mm, a maximum width of 8.6 mm, the rachis is 12.1 mm long, and the longest barb is 6.1 mm long. It lacks a plumulaceous portion. The barbs are equally spaced in a single plane, which indicates pennaceous vanes with intertwined barbules.
52. †Fossil feather 30 (UFRJ-DG 12 Av)
Early Cretaceous – CE
“UFRJ-DG 12 Av” – Metello 2017: 20, 24, 13.
A semiplume (but also classified as a contour feather in the same study) described by Metello ( 2017 ). It has a maximum length of 17.5 mm, a maximum width of 5.8 mm, the rachis is 13 mm long, and the longest barb is 11.6 mm long. The barbs are collapsed with the rachis. In general, the semiplumes he described have the rachis present as a thin dark line, as poorly preserved as the calami. The vanes have long and slender disarranged barbs, with non-intertwined barbules that show a fuzzy texture. The longest barbs are shorter than their respective rachises.
53. †Fossil feather 31 (UFRJ-DG 13 Av)
Early Cretaceous – CE
“UFRJ-DG 13 Av” – Metello 2017: 24.
A semiplume described by Metello ( 2017 ). It has a maximum length of 23 mm, a maximum width of 16 mm, the rachis is 19.7 mm long, and the longest barb is 13.8 mm long.
54. †Fossil feather 32 (UFRJ-DG 14 Av)
Early Cretaceous – CE
“UFRJ-DG 14 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 19.2 mm, a maximum width of 4.9 mm, the rachis is 16 mm long, and the longest barb is 10.2 mm long.
55. †Fossil feather 33 (UFRJ-DG 15 Av)
Early Cretaceous – CE
“UFRJ-DG 15 Av” – Metello 2017: 24.
A semiplume described by Metello ( 2017 ). It has a maximum length of 20.7 mm, a maximum width of 10.6 mm, the rachis is 14.9 mm long, and the longest barb is 10.4 mm long.
56. †Fossil feather 34 (UFRJ-DG 16 Av)
Early Cretaceous – CE
“UFRJ-DG 16 Av” – Metello 2017: 24, 14.
A semiplume described by Metello ( 2017 ). It has a maximum length of 18.9 mm, a maximum width of 13.2 mm, the rachis is 10.9 mm long, and the longest barb is 10.3 mm long.
57. †Fossil feather 35 (UFRJ-DG 17 Av)
Early Cretaceous – CE
“UFRJ-DG 17 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 10.6 mm, a maximum width of 7.6 mm, the rachis is 3.7 mm long, and the longest barb is 5.6 mm long.
58. †Fossil feather 36 (UFRJ-DG 18 Av)
Early Cretaceous – CE
“UFRJ-DG 18 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 17.3 mm, a maximum width of 4.8 mm, the rachis is 10.4 mm long, and the longest barb is 11.4 mm long.
59. †Fossil feather 37 (UFRJ-DG 19 Av)
Early Cretaceous – CE
“UFRJ-DG 19 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 16.5 mm, a maximum width of 14.6 mm, the rachis is 11.1 mm long, and the longest barb is 10.8 mm long.
60. †Fossil feather 38 (UFRJ-DG 20 Av)
Early Cretaceous – CE
“UFRJ-DG 20 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 14.15 mm, a maximum width of 8.7 mm, the rachis is 9.2 mm long, and the longest barb is 9.4 mm long.
61. †Fossil feather 39 (UFRJ-DG 21 Av)
Early Cretaceous – CE
“UFRJ-DG 21 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 17.2 mm, a maximum width of 10.2 mm, the rachis is 11.8 mm long, and the longest barb is 11.5 mm long.
62. †Fossil feather 40 (UFRJ-DG 22 Av)
Early Cretaceous – CE
“UFRJ-DG 22 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 13.5 mm, a maximum width of 4.7 mm, the rachis is 5.7 mm long, and the longest barb is 9.3 mm long.
63. †Fossil feather 41 (UFRJ-DG 23 Av)
Early Cretaceous – CE
“UFRJ-DG 23 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 20.8 mm, a maximum width of 10.1 mm, the rachis is 15.3 mm long, and the longest barb is 10.5 mm long.
64. †Fossil feather 42 (UFRJ-DG 24 Av)
Early Cretaceous – CE
“UFRJ-DG 24 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 16.2 mm, a maximum width of 8.1 mm, the rachis is 8.4 mm long, and the longest barb is 12.4 mm long.
65. †Fossil feather 43 (UFRJ-DG 25 Av)
Early Cretaceous – CE
“UFRJ-DG 25 Av” – Metello 2017: 24.
A semiplume described by Metello ( 2017 ). It has a maximum length of 15.8 mm, a maximum width of 7.4 mm, the rachis is 10.2 mm long, and the longest barb is 9.8 mm long.
66. †Fossil feather 44 (UFRJ-DG 26 Av)
Early Cretaceous – CE
“UFRJ-DG 26 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 10.7 mm, a maximum width of 6.6 mm, the rachis is 5 mm long, and the longest barb is 7.2 mm long.
67. †Fossil feather 45 (UFRJ-DG 27 Av)
Early Cretaceous – CE
“UFRJ-DG 27 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 15.8 mm, a maximum width of 10.9 mm, the rachis is 9.7 mm long, and the longest barb is 10.9 mm long.
68. †Fossil feather 46 (UFRJ-DG 28 Av)
Early Cretaceous – CE
“UFRJ-DG 28 Av” – Metello 2017: 24.
A semiplume described by Metello ( 2017 ). It has a maximum length of 15.9 mm, a maximum width of 9 mm, the rachis is 10.8 mm long, and the longest barb is 9.1 mm long.
69. †Fossil feather 47 (UFRJ-DG 29 Av)
Early Cretaceous – CE
“UFRJ-DG 29 Av” – Metello 2017: 24.
A semiplume described by Metello ( 2017 ). It has a maximum length of 20 mm, a maximum width of 17.5 mm, the rachis is 15.1 mm long, and the longest barb is 10.7 mm long.
70. †Fossil feather 48 (UFRJ-DG 32 Av)
Early Cretaceous – CE
“UFRJ-DG 32 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 20.2 mm, a maximum width of 9.8 mm, the rachis is 8.5 mm long, and the longest barb is 14.6 mm long.
71. †Fossil feather 49 (UFRJ-DG 33 Av)
Early Cretaceous – CE
“UFRJ-DG 33 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 31.7 mm, a maximum width of 4.5 mm, the rachis is about 16.9 mm long, and the longest barb is 9.12 mm long.
72. †Fossil feather 50 (UFRJ-DG 35 Av)
Early Cretaceous – CE
“UFRJ-DG 35 Av” – Metello 2017: 19–20, 31, 12.
A contour feather described by Metello ( 2017 ). It has a maximum length of 19.4 mm, a maximum width of 14.2 mm, the rachis is 12.9 mm long, and the longest barb is 12 mm long. It shows open and loose vane edges and was possibly used for sexual display.
73. †Fossil feather 51 (UFRJ-DG 36 Av)
Early Cretaceous – CE
“UFRJ-DG 36 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 18.1 mm, a maximum width of 7.6 mm, the rachis is 10.2 mm long, and the longest barb is 10.8 mm long.
74. †Fossil feather 52 (UFRJ-DG 37 Av)
Early Cretaceous – CE
“UFRJ-DG 37 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). It has a maximum length of 15.5 mm, a maximum width of 7.5 mm, the rachis is 9.1 mm long, and the longest barb is 8.9 mm long. It is curved, with loose barbs at the feather edge, similar to wing coverts of modern birds.
75. †Fossil feather 53 (UFRJ-DG 38 Av)
Early Cretaceous – CE
“UFRJ-DG 38 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 21.3 mm, a maximum width of 18.2 mm, the rachis is 13.4 mm long, and the longest barb is 14.3 mm long.
76. †Fossil feather 54 (UFRJ-DG 39 Av)
Early Cretaceous – CE
“UFRJ-DG 39 Av” – Metello 2017: 19–20, 11.
A contour feather described by Metello ( 2017 ). It has a maximum length of 21.6 mm, a maximum width of 15.3 mm, the rachis is 15.9 mm long, and the longest barb is 11.6 mm long. The barbs are collapsed with the rachis. The plumulaceous area of the feather accounts for about half of the total length of the vanes.
77. †Fossil feather 55 (UFRJ-DG 40 Av)
Early Cretaceous – CE
“UFRJ-DG 40 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). It has a maximum length of 25.9 mm, a maximum width of 6.8 mm, the rachis is 18.2 mm long, and the longest barb is 12 mm long. The barbs are collapsed with the rachis.
78. †Fossil feather 56 (UFRJ-DG 41 Av)
Early Cretaceous – CE
“UFRJ-DG 41 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 15.8 mm, a maximum width of 6.5 mm, the rachis is 8.9 mm long, and the longest barb is 6.3 mm long.
79. †Fossil feather 57 (UFRJ-DG 42 Av)
Early Cretaceous – CE
“UFRJ-DG 42 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 15.5 mm, a maximum width of 15.2 mm, the rachis is 11.2 mm long, and the longest barb is 11.3 mm long.
80. †Fossil feather 58 (UFRJ-DG 43 Av)
Early Cretaceous – CE
“UFRJ-DG 43 Av” – Metello 2017: 20, 24.
A semiplume (but also suggested to be a contour feather in the same study) described by Metello ( 2017 ). It has a maximum length of 14.1 mm, a maximum width of 4.85 mm, the rachis is 8.9 mm long, and the longest barb is 6.7 mm long. The barbs are collapsed with the rachis.
81. †Fossil feather 59 (UFRJ-DG 44 Av)
Early Cretaceous – CE
“UFRJ-DG 44 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). The barbs are equally spaced in a single plane, which indicates pennaceous vanes with intertwined barbules. The longest barb is 6.9 mm long.
82. †Fossil feather 60 (UFRJ-DG 45 Av)
Early Cretaceous – CE
“UFRJ-DG 45 Av” – Metello 2017: 24.
A plume described by Metello ( 2017 ). It has a maximum length of 9.1 mm, a maximum width of 6.9 mm, the rachis is 4.8 mm long, and the longest barb is 6.6 mm long.
83. †Fossil feather 61 (UFRJ-DG 46 Av)
Early Cretaceous – CE
“UFRJ-DG 46 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). It has a maximum length of 16.7 mm, a maximum width of 7.9 mm, the rachis is 11.9 mm long, and the longest barb is 9.3 mm long. The barbs are equally spaced in a single plane, which indicates pennaceous vanes with intertwined barbules.
84. †Fossil feather 62 (UFRJ-DG 47 Av)
Early Cretaceous – CE
“UFRJ-DG 47 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 23.5 mm, a maximum width of 17.5 mm, the rachis is 15.8 mm long, and the longest barb is 7.5 mm long.
85. †Fossil feather 63 (UFRJ-DG 48 Av)
Early Cretaceous – CE
“UFRJ-DG 48 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 19.3 mm, a maximum width of 6.8 mm, the rachis is 14.9 mm long, and the longest barb is 10.3 mm long.
86. †Fossil feather 64 (UFRJ-DG 49 Av)
Early Cretaceous – CE
“UFRJ-DG 49 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). It has a maximum length of 19.9 mm, a maximum width of 5.7 mm, the rachis is 13.4 mm long, and the longest barb is 11.9 mm long. The barbs are collapsed with the rachis.
87. †Fossil feather 65 (UFRJ-DG 50 Av)
Early Cretaceous – CE
“UFRJ-DG 50 Av” – Metello 2017: 19–20.
A contour feather described by Metello ( 2017 ). It has a maximum length of 19.1 mm, a maximum width of 4.5 mm, the rachis is 10.3 mm long, and the longest barb is 14.4 mm long. It is curved, with loose barbs at the feather edge, similar to wing coverts of modern birds.
88. †Fossil feather 66 (UFRJ-DG 51 Av)
Early Cretaceous – CE
“UFRJ-DG 51 Av” – Metello 2017: 24.
A semiplume described by Metello ( 2017 ). It has a maximum length of 21.5 mm, a maximum width of 8.5 mm, the rachis is 18 mm long, and the longest barb is 13.1 mm long.
89. †Fossil feather 67 (UFRJ-DG 52 Av)
Early Cretaceous – CE
“UFRJ-DG 52 Av” – Metello 2017: 24, 15.
A plume described by Metello ( 2017 ). It has a maximum length of 14.7 mm, a maximum width of 11.9 mm, the rachis is 7.4 mm long, and the longest barb is 8.7 mm long.
90. †Fossil feather 68 (UFRJ-DG 53 Av)
Early Cretaceous – CE
“UFRJ-DG 53 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 17.1 mm, a maximum width of 4.8 mm, the rachis is 10.3 mm long, and the longest barb is 10.1 mm long.
91. †Fossil feather 69 (UFRJ-DG 54 Av)
Early Cretaceous – CE
“UFRJ-DG 54 Av” – Metello 2017: 19.
A contour feather described by Metello ( 2017 ). It has a maximum length of 13.7 mm, a maximum width of 6.4 mm, the rachis is 8.7 mm long, and the longest barb is 8.2 mm long.
92. †Fossil feather 70 (CAV-003-V)
Early Cretaceous – CE
“CAV-003-V” – França et al. 2017: 19.
França et al. ( 2017 ) reported this plumaceous feather from the Crato Formation. It has a rachis 11.35 mm long and a barb 6.58 mm long. The calamus is absent.
93. †Fossil feather 71 (CAV-004-V)
Early Cretaceous – CE
“CAV-004-V” – França et al. 2017: 19.
França et al. ( 2017 ) reported this plumaceous feather from the Crato Formation. It has a rachis 15.88 mm long and its largest barb is 8.95 mm long. The calamus is absent.
94. †Fossil feather 72 (LPU-271)
Early Cretaceous – CE
“LPU-271” – França et al. 2017: 19.
França et al. ( 2017 ) reported this plumaceous feather from the Crato Formation. It has a rachis 9.74 mm long and its largest barb is 6.1 mm long. The calamus is absent.
95. †Fossil feather 73 (LPU-520a)
Early Cretaceous – CE
“LPU-520a” – França et al. 2017: 19.
França et al. ( 2017 ) reported this plumaceous feather from the Crato Formation. It has a rachis 16.05 mm long, calamus 1.33 mm long and a barb 7.02 mm long.
96. †Fossil feather 74 (LPU-1464b)
Early Cretaceous – CE
“LPU-1464b” – França et al. 2017: 19.
França et al. ( 2017 ) reported this plumaceous feather from the Crato Formation. It has a rachis 19.24 mm long and its largest barb is 9.47 mm long. The calamus is absent.
97. †Fossil feather 75 (LPU-272)
Early Cretaceous – CE
“LPU-272” – Lopes et al. 2017: 20.
Lopes et al. ( 2017 ) reported this contour feather from the Crato Formation. It is 18.48 mm long.
98. †Fossil feather 76 (LPU-521)
Early Cretaceous – CE
“LPU-521” – Lopes et al. 2017: 20.
Lopes et al. ( 2017 ) reported this contour feather from the Crato Formation.
99. †Fossil feather 77 (LPU-522)
Early Cretaceous – CE
“LPU-522” – Lopes et al. 2017: 20.
Lopes et al. ( 2017 ) reported this contour feather from the Crato Formation. It is 9.01 mm long.
100. †Fossil feather 78 (LPU-523)
Early Cretaceous – CE
“LPU-523” – Lopes et al. 2017: 20.
Lopes et al. ( 2017 ) reported this contour feather from the Crato Formation.
101. †Fossil feather 79 (MPSC-PN 1117)
Nascimento ( 2020 ) reported this well-preserved plume from the Crato Formation. It is 6.95 mm wide and 15.25 mm long. The rachis, barbs, barbules, and calamus are present.
102. †Fossil feather 80 (MPSC-PN 1233)
Nascimento ( 2020 ) reported this well-preserved semiplume from the Crato Formation. It is 6.55 mm wide and 15.69 mm long. The rachis and barbs are present, but barbules are not visible.
103. †Fossil feather 81 (MPSC-PN 1286 AB)
Nascimento ( 2020 ) reported this semiplume as well-preserved in part and counterpart from the Crato Formation. It is 6.61 mm wide and 12.65 mm long. The rachis, barbs, barbules, and a partial calamus are present.
104. †Fossil feather 82 (MPSC-PN 2382)
Nascimento ( 2020 ) reported this contour feather from the Crato Formation. It is 5.7 mm wide and 19.4 mm long. The rachis, barbs, and barbules are well-preserved. A banded pattern of dark and light stripes is also present.
105. †Fossil feather 83 (MPSC-PN 2383)
Nascimento ( 2020 ) reported this well-preserved plume from the Crato Formation. It is 9.28 mm wide and 9.72 mm long. The rachis, barbs, barbules, and a partial calamus are present.
106. †Fossil feather 84 (UFRPE 5019)
Nascimento ( 2020 ) reported this well-preserved contour feather from the Crato Formation. It is 7.24 mm wide and 14.86 mm long. The rachis, barbs, and barbules are present.
107. †Fossil feather 85 (UFRPE 5113 AB)
Nascimento ( 2020 ) reported this contour feather preserved in part and counterpart from the Crato Formation. It is 14.44 mm wide and 19.96 mm long. The rachis, barbs, and barbules are well preserved.
108. †Fossil feather 86 (“URCA 1 AB”)
Nascimento ( 2020 ) reported this contour feather preserved in part and counterpart from the Crato Formation. It is 9.25 mm wide and 18.5 mm long. The rachis, barbs, barbules, and calamus are present. This specimen and the following six ones received a provisional museum number since they were not cataloged yet.
109. †Fossil feather 87 (“URCA 2 AB”)
Nascimento ( 2020 ) reported this plume preserved in part and counterpart from the Crato Formation. It is 13.2 mm wide and 13.2 mm long. The rachis, barbs, barbules, and calamus are present.
110. †Fossil feather 88 (“URCA 3”)
Nascimento ( 2020 ) reported this well-preserved plume from the Crato Formation. It is 5.98 mm wide and 7.85 mm long. The rachis, barbs, barbules, and calamus are present.
111. †Fossil feather 89 (“URCA 4 AB”)
Nascimento ( 2020 ) reported this semiplume preserved in part and counterpart from the Crato Formation. It is 4.82 mm wide and 22.85 mm long. One of the parts holds most of the feather impression. Parts of the rachis, barbs, and barbules are visible despite the poor preservation and inadequate preparation.
112. †Fossil feather 90 (“URCA 5”)
Nascimento ( 2020 ) reported this contour feather from the Crato Formation. It is 7.06 mm wide and 17.34 mm long. The rachis, barbs, and barbules are visible.
113. †Fossil feather 91 (“URCA 6”)
Nascimento ( 2020 ) reported this contour feather preserved in part and counterpart from the Crato Formation. It is 8.93 mm wide and 19.38 mm long. One part holds most of the feather, while the other has only a very faint impression. The rachis, barbs, and barbules are visible. The calamus might become visible if the specimen goes through a more complete preparation. A banded pattern of dark and light stripes is also present.
114. †Fossil feather 92 (“URCA 7 AB”)
Nascimento ( 2020 ) reported this plume preserved in part and counterpart from the Crato Formation. It is 9.11 mm wide and 15.38 mm long. The rachis, barbs, and barbules are visible. The calamus might become visible if the specimen goes through a more complete preparation.
115. †Fossil feather 93
Early Cretaceous – CE
“pena (ave?)” – Bantim et al. 2022: 114.
Bantim et al. ( 2022 ) mentioned a feather among fossil material from the Crato Formation handed by miners as part of an educational and awareness program created in 2021 by the Araripe GeoPark Mundial of UNESCO, MPSC (where the material is deposited), and Batalhão de Polícia do Meio Ambiente.
116. †Fossil feather 94
Early Cretaceous – CE
“Penas não-avianas” [in part] – Sousa et al. ( 2023 ): 79.
Sousa et al. ( 2023 ) reported two contour feathers from the Crato Formation of Santana do Cariri. They were collected in 2015 by professors of UESB and are housed in the Acervo do Laboratório de Geociências II (LabGeoc) of that institution. The first of them is 10 mm long and preserved the rachis, barbs, and barbules. Although the authors described the feathers as non-avian, no argument was presented to support this.
117. †Fossil feather 95
Early Cretaceous – CE
“Penas não-avianas” [in part] – Sousa et al. ( 2023 ): 79.
The second contour feather reported by Sousa et al. ( 2023 ) is 11 mm long and preserved a small rachis, barbs, barbules, and calamus.
118. †Fossil feather 96 (GP/2E-9378 and GP/2E-9381)
Early Cretaceous – CE
Fig. 7 C–D
A dark-colored feather preserved in part and counterpart about 20 mm long from the Crato Formation. It lacks a visible calamus.
119. †Fossil feather 97 (GP/2E-9379 and GP/2E-9405)
Early Cretaceous – CE
Fig. 7 F–G
A dark-colored feather preserved in part and counterpart about 19 mm long from the Crato Formation. It lacks a visible calamus.
120. †Fossil feather 98 (GP/2E-9380)
Early Cretaceous – CE
Fig. 7 H
A light-colored feather about 29 mm long from the Crato Formation.
121. †Fossil feather 99 (GP/2E-9382)
Early Cretaceous – CE
Fig. 7 I
A dark-colored feather about 13 mm long from the Crato Formation. It lacks a visible calamus.
122. †Fossil feather 100 (GP/2E-9383)
Early Cretaceous – CE
Fig. 7 J
A dark-colored feather about 13 mm long from the Crato Formation. It lacks a visible calamus.
123. †Fossil feather 101 (GP/2E-9384 and GP/2E-9428)
Early Cretaceous – CE
Fig. 7 K–L
A dark-colored feather preserved in part and counterpart about 20 mm long from the Crato Formation. It lacks a visible calamus.
124. †Fossil feather 102 (GP/2E-9385)
Early Cretaceous – CE
Fig. 7 M
A dark-colored feather about 18 mm long from the Crato Formation. It lacks a visible calamus.
125. †Fossil feather 103 (GP/2E-9386)
Early Cretaceous – CE
Fig. 7 N
A faintly preserved feather about 17 mm long from the Crato Formation. It lacks a visible calamus.
126. †Fossil feather 104 (GP/2E-9387 and GP/2E-9406)
Early Cretaceous – CE
Fig. 7 O–P
A dark-colored feather well-preserved in part and counterpart about 13 mm long from the Crato Formation. It lacks a visible calamus.
127. †Fossil feather 105 (GP/2E-9388 and GP/2E-9416)
Early Cretaceous – CE
Fig. 8 C–D
A dark-colored feather poorly preserved in part and counterpart about 13 mm long from the Crato Formation. It lacks a visible calamus.
128. †Fossil feather 106 (GP/2E-9389)
Early Cretaceous – CE
Fig. 8 A
A possible flight feather, well-preserved, light-colored, about 49 mm long from the Crato Formation.
129. †Fossil feather 107 (GP/2E-9390)
Early Cretaceous – CE
Fig. 8 B
A possible tail feather, well-preserved, light-colored, about 40 mm long from the Crato Formation.
130. †Fossil feather 108 (GP/2E-9391)
Early Cretaceous – CE
Fig. 8 E
A well-preserved dark-colored feather about 18 mm long from the Crato Formation. It lacks a visible calamus.
131. †Fossil feather 109 (GP/2E-9392)
Early Cretaceous – CE
Fig. 8 F
A poorly preserved dark-colored feather about 19 mm long from the Crato Formation. It lacks a visible calamus.
132. †Fossil feather 110 (GP/2E-9393)
Early Cretaceous – CE
Fig. 8 G
A mildly preserved dark-colored feather about 14 mm long from the Crato Formation. It lacks a visible calamus.
133. †Fossil feather 111 (GP/2E-9394)
Early Cretaceous – CE
Fig. 8 H
A mildly preserved light-colored feather about 14 mm long from the Crato Formation. It lacks a visible calamus.
134. †Fossil feather 112 (GP/2E-9395)
Early Cretaceous – CE
Fig. 8 I
A well-preserved dark-colored feather about 12 mm long from the Crato Formation. It lacks a visible calamus.
135. †Fossil feather 113 (GP/2E-9396 and GP/2E-9424)
Early Cretaceous – CE
Fig. 8 K–L
A dark-colored feather poorly preserved in part and counterpart about 15 mm long from the Crato Formation. It lacks a visible calamus.
136. †Fossil feather 114 (GP/2E-9397)
Early Cretaceous – CE
Fig. 8 J
A poorly preserved light-colored feather about 24 mm long from the Crato Formation.
137. †Fossil feather 115 (GP/2E-9398)
Early Cretaceous – CE
Fig. 8 M
A well-preserved dark-colored feather about 13 mm long from the Crato Formation. It lacks a visible calamus.
138. †Fossil feather 116 (GP/2E-9399)
Early Cretaceous – CE
Fig. 8 N
A well-preserved dark-colored feather about 17 mm long from the Crato Formation. It lacks a visible calamus.
139. †Fossil feather 117 (GP/2E-9400)
Early Cretaceous – CE
Fig. 9 A
A well-preserved dark-colored feather about 17 mm long from the Crato Formation. It lacks a visible calamus.
140. †Fossil feather 118 (GP/2E-9401)
Early Cretaceous – CE
Fig. 9 B
A mildly preserved dark-colored feather about 14 mm long from the Crato Formation.
141. †Fossil feather 119 (GP/2E-9402 and GP/2E-9437)
Early Cretaceous – CE
Fig. 9 C–D
A dark-colored feather mildly preserved in part and counterpart about 16 mm long from the Crato Formation. It lacks a visible calamus.
142. †Fossil feather 120 (GP/2E-9403)
Early Cretaceous – CE
Fig. 9 E
A poorly preserved dark-colored feather about 15 mm long from the Crato Formation. It lacks a visible calamus.
143. †Fossil feather 121 (GP/2E-9404 and GP/2E-9417)
Early Cretaceous – CE
Fig. 9 F–G
A mildly dark-colored feather preserved in part and counterpart about 13 mm long from the Crato Formation. It lacks a visible calamus.
144. †Fossil feather 122 (GP/2E-9407)
Early Cretaceous – CE
Fig. 9 H
A poorly preserved dark-colored feather about 21 mm long from the Crato Formation. It lacks a visible calamus.
145. †Fossil feather 123 (GP/2E-9408)
Early Cretaceous – CE
Fig. 9 I
A well-preserved, light-colored feather about 11 mm long from the Crato Formation. It lacks a visible calamus.
Feathers from the Crato Formation: GP/2E-9389 (A); GP/2E-9390 (B); GP/2E-9388 (C); GP/2E-9416 (D); GP/2E-9391 (E); GP/2E-9392 (F); GP/2E-9393 (G); GP/2E-9394 (H); GP/2E-9395 (I); GP/2E-9397 (J); GP/2E-9396 (K); GP/2E-9424 (L); GP/2E-9398 (M); GP/2E-9399 (N) (R.S. Nascimento). Scale bar: 5 mm.
146. †Fossil feather 124 (GP/2E-9409)
Early Cretaceous – CE
Fig. 9 J
A poorly preserved dark-colored feather about 12 mm long from the Crato Formation. It lacks a visible calamus.
147. †Fossil feather 125 (GP/2E-9410 and GP/2E-9415)
Early Cretaceous – CE
Fig. 9 K–L
A dark-colored feather well-preserved in part and counterpart about 19 mm long from the Crato Formation. It lacks a visible calamus.
148. †Fossil feather 126 (GP/2E-9411)
Early Cretaceous – CE
Fig. 9 M
A poorly preserved dark-colored feather about 16 mm long from the Crato Formation. It lacks a visible calamus.
149. †Fossil feather 127 (GP/2E-9412)
Early Cretaceous – CE
Fig. 9 N
A poorly preserved dark-colored feather about 15 mm long from the Crato Formation. It lacks a visible calamus.
150. †Fossil feather 128 (GP/2E-9413)
Early Cretaceous – CE
Fig. 9 O
A mildly preserved dark-colored feather about 14 mm long from the Crato Formation. It lacks a visible calamus.
151. †Fossil feather 129 (GP/2E-9414)
Early Cretaceous – CE
Fig. 9 P
A poorly preserved dark-colored feather about 19 mm long from the Crato Formation. It lacks a visible calamus.
152. †Fossil feather 130 (GP/2E-9418)
Early Cretaceous – CE
Fig. 10 A
A mildly preserved dark-colored feather about 15 mm long from the Crato Formation. It lacks a visible calamus.
153. †Fossil feather 131 (GP/2E-9419)
Early Cretaceous – CE
Fig. 10 B
A poorly preserved dark-colored feather about 20 mm long from the Crato Formation. It lacks a visible calamus.
154. †Fossil feather 132 (GP/2E-9420)
Early Cretaceous – CE
Fig. 10 C
A poorly preserved dark-colored feather about 15 mm long from the Crato Formation. It lacks a visible calamus.
155. †Fossil feather 133 (GP/2E-9421)
Early Cretaceous – CE
Fig. 10 D
A mildly preserved dark-colored feather about 10 mm long from the Crato Formation. It lacks a visible calamus.
156. †Fossil feather 134 (GP/2E-9422 and GP/2E-9442)
Early Cretaceous – CE
Fig. 10 E–F
A dark-colored feather poorly preserved in part and counterpart about 26 mm long from the Crato Formation. It lacks a visible calamus.
157. †Fossil feather 135 (GP/2E-9423)
Early Cretaceous – CE
Fig. 10 G
A poorly preserved dark-colored feather about 18 mm long from the Crato Formation. It lacks a visible calamus.
158. †Fossil feather 136 (GP/2E-9425)
Early Cretaceous – CE
Fig. 10 H
A mildly preserved proximally dark- and distally light-colored feather about 10 mm long from the Crato Formation.
159. †Fossil feather 137 (GP/2E-9426)
Early Cretaceous – CE
Fig. 10 I
A well-preserved dark-colored feather about 15 mm long from the Crato Formation.
160. †Fossil feather 138 (GP/2E-9427 and GP/2E-9440)
Early Cretaceous – CE
Fig. 10 J–K
A light-colored feather mildly preserved in part and counterpart about 16 mm long from the Crato Formation. It lacks a visible calamus.
161. †Fossil feather 139 (GP/2E-9429)
Early Cretaceous – CE
Fig. 10 L
A well-preserved, incompletely collected dark-colored feather with a preserved middle to distal length of about 18 mm from the Crato Formation.
Feathers from the Crato Formation: GP/2E-9400 (A); GP/2E-9401 (B); GP/2E-9402 (C); GP/2E-9437 (D); GP/2E-9403 (E); GP/2E-9404 (F); GP/2E-9417 (G); GP/2E-9407 (H); GP/2E-9408 (I); GP/2E-9409 (J); GP/2E-9410 (K); GP/2E-9415 (L); GP/2E-9411 (M); GP/2E-9412 (N); GP/2E-9413 (O); GP/2E-9414 (P) (R.S. Nascimento). Scale bar: 5 mm.
162. †Fossil feather 140 (GP/2E-9430)
Early Cretaceous – CE
Fig. 10 M
A faintly preserved dark-colored feather about 20 mm long from the Crato Formation. It lacks a visible calamus.
163. †Fossil feather 141 (GP/2E-9431)
Early Cretaceous – CE
Fig. 10 N
A mildly preserved dark-colored feather about 10 mm long from the Crato Formation. It lacks a visible calamus.
164. †Fossil feather 142 (GP/2E-9432)
Early Cretaceous – CE
Fig. 10 O
A well-preserved dark-colored feather about 17 mm long from the Crato Formation.
165. †Fossil feather 143 (GP/2E-9433)
Early Cretaceous – CE
Fig. 10 P
A mildly preserved dark-colored feather about 12 mm long from the Crato Formation. It lacks a visible calamus.
166. †Fossil feather 144 (GP/2E-9434)
Early Cretaceous – CE
Fig. 11 A
A mildly preserved dark-colored feather about 7 mm long from the Crato Formation. It lacks a visible calamus.
167. †Fossil feather 145 (GP/2E-9435)
Early Cretaceous – CE
Fig. 11 B
A poorly preserved dark-colored feather about 14 mm long from the Crato Formation.
168. †Fossil feather 146 (GP/2E-9436)
Early Cretaceous – CE
Fig. 11 C
A poorly preserved dark-colored feather about 27 mm long from the Crato Formation. It lacks the proximal part.
169. †Fossil feather 147 (GP/2E-9438)
Early Cretaceous – CE
Fig. 11 D
A well-preserved dark-colored feather about 13 mm long from the Crato Formation. It lacks a visible calamus.
170. †Fossil feather 148 (GP/2E-9439)
Early Cretaceous – CE
Fig. 11 E
A poorly preserved dark-colored feather about 14 mm long from the Crato Formation. It lacks a visible calamus.
171. †Fossil feather 149 (GP/2E-9441)
Early Cretaceous – CE
Fig. 11 F
A mildly preserved dark-colored feather about 14 mm long from the Crato Formation.
172. †Fossil feather 150 (GP/2E-9443)
Early Cretaceous – CE
Fig. 11 G
A faintly preserved light-colored feather about 22 mm long from the Crato Formation.
173. †Fossil feather 151 (GP/2E-9467)
Early Cretaceous – CE
Fig. 11 H
A mildly preserved dark-colored feather about 19 mm long from the Crato Formation.
174. †Fossil feather 152 (GP/2E-9476 and GP/2E-9477)
Early Cretaceous – CE
Fig. 11 I–J
A dark-colored feather poorly preserved in part and counterpart about 16 mm long from the Crato Formation. It lacks a visible calamus.
175. †Fossil feather 153 (GP/2E-9478 and GP/2E-9479)
Early Cretaceous – CE
Fig. 11 K–L
A dark-colored feather faintly preserved in part and counterpart about 21 mm long from the Crato Formation. It lacks a visible calamus.
176. †Fossil feathers 154 (GP/2E-9480 and GP/2E-9481)
Early Cretaceous – CE
Fig. 11 M–N
Dark-colored, entangled feathers (apparently three) from the Crato Formation, preserved as part and counterpart.
177. †Fossil feather 155 (GP/2E-9482 and GP/2E-9483)
Early Cretaceous – CE
Fig. 11 O–P
A dark-colored feather about 18 mm long poorly preserved in part and counterpart from the Crato Formation. It lacks a visible calamus.
Feathers from the Crato Formation: GP/2E-9418 (A); GP/2E-9419 (B); GP/2E-9420 (C); GP/2E-9421 (D); GP/2E-9422 (E); GP/2E-9442 (F); GP/2E-9423 (G); GP/2E-9425 (H); GP/2E-9426 (I); GP/2E-9427 (J); GP/2E-9440 (K); GP/2E-9429 (L); GP/2E-9430 (M); GP/2E-9431 (N); GP/2E-9432 (O); GP/2E-9433 (P) (R.S. Nascimento). Scale bar: 5 mm.
Feathers from the Crato Formation: GP/2E-9434 (A); GP/2E-9435 (B); GP/2E-9436 (C); GP/2E-9438 (D); GP/2E-9439 (E); GP/2E-9441 (F); GP/2E-9443 (G); GP/2E-9467 (H); GP/2E-9476 (I); GP/2E-9477 (J); GP/2E-9478 (K); GP/2E-9479 (L); GP/2E-9480 (M); GP/2E-9481 (N); GP/2E-9482 (O); GP/2E-9483 (P) (R.S. Nascimento). Scale bar: 5 mm.
178. †Fossil feather 156 (GP/2E-9484)
Early Cretaceous – CE
Fig. 12 A
A well-preserved dark-colored feather about 18 mm long from the Crato Formation.
179. †Fossil feather 157 (GP/2E-9485)
Early Cretaceous – CE
Fig. 12 B
A mildly preserved dark-colored feather about 12 mm long from the Crato Formation. It lacks a visible calamus.
180. †Fossil feather 158 (GP/2E-9486)
Early Cretaceous – CE
Fig. 12 C
A well-preserved dark-colored feather about 14 mm long from the Crato Formation. It lacks a visible calamus.
181. †Fossil feather 159 (GP/2E-9494)
Early Cretaceous – CE
Fig. 12 D
A well-preserved dark-colored feather about 19 mm long from the Crato Formation. It lacks a visible calamus.
182. †Fossil feather 160 (LEG 0766 and LE G0896)
Early Cretaceous – CE
Fig. 12 E–F
A possible flight feather preserved in part and counterpart from the Pedra Cariri site in Nova Olinda, Ceará.
183. †Fossil feather 161 (LEG 0885 and LEG 0886)
Early Cretaceous – CE
Fig. 12 G–H
A possible flight feather preserved in part and counterpart from the Pedra Cariri site in Nova Olinda, Ceará. It preserves a possible color pattern.
184. †Fossil feather 162 (MHNCE/FOS-001)
Early Cretaceous – CE
Fig. 12 I
A mildly preserved dark-colored feather about 23 mm long from the Crato Formation. It lacks a visible calamus.
2.4.4.2. Cenozoic feathers
185. † Fossil feathers 163 (Entre-Córregos Formation)
The occurrence of feathers in the Entre-Córregos Formation was first reported by Bedani and Haddad ( 2002 ). Subsequent authors (e.g., Franco-Delgado and Bernardes-de-Oliveira 2004 , Martins Neto and Pesenti 2006a , 2006b , Castro-Fernandes et al. 2013 ) also mentioned them, but without a proper description or figures.
186. †Fossil feather 164 (Museu Paulista 111)
Late Oligocene/Early Miocene – SP
“Fossil feather” – Shufeldt 1916: 206–207.
Aves [“Indeterminadas”; in part] – Paula Couto 1958a: 2, 5.
Aves [“Indeterminadas”; in part] – Paula Couto 1970: 922.
“penas fósseis” [in part] – Sick 1984a: 65.
“fossil feathers” [in part] – Alvarenga 1993a: 62.
“penas fósseis” [in part] – Alvarenga 1993b: 21.
“penas fósseis” [in part] – Alvarenga 1997: 123.
“penas isoladas” [in part] – Kellner 1998: 659.
“primary feather of a large bird” – Sayão et al. 2011: 198.
“One primary remex” – Prado et al. 2016a: 4.
Shufeldt ( 1916 ) reported a feather from the shales of the Tremembé Formation. The specimen, No. 111 at the Museu Paulista, was sent to him by the institution’s director, Dr. von Ihering, in a letter of transmission dated 8 January 1915.
The fossil matrix has a dark chocolate-brown color, with roughness similar to leather on the face on which the feather is found, whereas the posterior face is lighter in color, with evidence of horizontal cleavage. The plate is 14 x 7.5 cm and has an average thickness of 3 mm, with markings that indicate it was collected with some sharp instrument, such as a knife. When the plate is wet, the feather, which is quite faint, becomes more evident. The feather was preserved as a carbonized trace (Prado et al. 2016a ). It has a total length of 113 mm, the calamus is 40 mm long, and the vane is 73 mm long. Despite its plumular appearance, Shufeldt believed it to be a primary feather of a sizable bird.
Shufeldt took a life-size photograph of the specimen to archive with similar ones in his collection (but this has not been published). He noted it is impossible to perceive the feather impressions smallest structures, even with a powerful lens. Although Silva Santos ( 1950 ) reiterated that the specimen was at the time in the Museu Paulista (probably only reproducing what Shufeldt published), when, or whether, the fossil was returned is unknown. The zoological and paleontological collections of the Museu Paulista were eventually transferred to the MZUSP. The fossil feather, however, could not be located.
187. †Fossil feather 165 (DGM 1-A)
Late Oligocene/Early Miocene – SP
Fig. 12 J–K
“uma das penas de cobertura do corpo-pluma de um passeriforme” – Silva Santos 1950: 445–446, fig. in pp. 445.
Aves [“Indeterminadas”; in part] – Paula Couto 1958a: 2, 5.
“Turdídeo (?)” – Mezzalira 1966: 102, pl. IX 4.
Aves [“Indeterminadas”; in part] – Paula Couto 1970: 922.
“penas fósseis” [in part] – Sick 1984a: 65.
“Pena de ave não identificada” – Lima 1989: 76, fig. in pp. 76.
“Turdídeo (?)” – Mezzalira 1989: 111, pl. IX 4.
“fossil feathers” [in part] – Alvarenga 1993a: 62.
“penas fósseis” [in part] – Alvarenga 1993b: 21.
“penas fósseis” [in part] – Alvarenga 1997: 123.
“penas isoladas” [in part] – Kellner 1998: 659.
“isolated feathers imprints” [sic] – Sayão et al. 2011: 198.
“One contour feather” – Prado et al. 2016a: 4.
A second feather from the Tremembé Formation was described by Silva Santos ( 1950 ). Gilberto H. William collected the material (originally DGM 1-A, pending new MCT code) in July 1948 at the Mina Nossa Senhora da Guia in a layer of dark green shales approximately 12.5 m deep, during the paleontological research carried out by DGM of DNPM.
It was preserved as part and counterpart. Silva Santos described and depicted the longest carbonized impression (27 mm) as a contour feather of a passeriform, of size close to a turdid. The calamus is absent, but when considering the average length for such a structure, the complete feather should have around 30 mm. Silva Santos also noted that, although the specimen appears very well preserved to the naked eye, no study under a microscope could be done on its minute structure, thus preventing an attempt at classification.
188. †Fossil feather 166 (Tremembé Formation)
Late Oligocene/Early Miocene – SP
“impressões de penas” – Garcia 1993: 329.
Garcia ( 1993 ) reported feathers from the Tremembé Formation in the UNG collection, among the material from the deposits of Fazenda Santa Fé, Mineração Aligra S/A, and Nossa Senhora da Guia, in Tremembé and Taubaté.
189. †Fossil feather 167 (GP/2E-8125)
Prado and Anelli ( 2014a ) reported a contour feather (GP/2E-8125) from the Tremembé Formation obtained in 2006 at the Sociedade Extrativa Santa Fé quarry. Prado and Anelli ( 2015 ) performed SEM analysis on the specimen and the observed microbodies were interpreted as eumelanosomes, which suggests it had a dark coloration. Prado et al. ( 2016b ) described it as an incomplete contour feather composed only by the mid portion, presenting a black hue, with preserved barbs and barbules. It is 17.17 mm wide and 26.42 mm long. The largest barb is 19.62 mm, and the smallest is 1.88 mm long. The rachis is not visible. Ostracod shells were preserved in the matrix. Prado ( 2017 ) reported that the eumelanosomes’ morphology in this specimen suggests it was dark or slightly dark-colored, possibly grayish.
190. †Fossil feather 168 (GP/2E-8126 and GP/2E-8127)
Prado and Anelli ( 2014a ) reported an incomplete rectrix discovered in 2006 in the Sociedade Extrativa Santa Fé quarry as two separate specimens (GP/2E-8126 and GP/2E-8127). Prado et al. ( 2016b ) reported it as a single feather preserved in part and counterpart with identifiable barbs and a thick rachis. The GP/2E-8126 specimen is 23.65 mm wide and 67.90 mm long, its largest barb is 19.47 mm, and the smallest is 4.21 mm long, and its rachis is 64.74 mm long and 1.36 mm thick. The GP/2E-8127 specimen is 32.33 mm wide and 52.33 mm long, its largest barb is 26.67 mm, and the smallest is 3.33 mm long, and its rachis is 52.33 mm long and 3.67 mm thick. The rachis and the vanes appear to have been severely crushed and chemically altered. A small fish vertebra is present in the surrounding matrix. The authors suggested it possibly belonged to aquatic taxa considering its taphonomic history. Its size indicates a medium-sized bird not longer than 80–90 cm.
191. †Fossil feather 169 (Pirabas Formation)
Ackermann ( 1964 ) reported a feather preserved as a carbonized trace in limestone from the Caieira site (Olaria) of Pirabas Formation ( Fig. 1 .8) (early Miocene), near Capanema, Pará (Távora et al. 2010 , Prado et al. 2016a ). He noted that the fossil “still retains the beautiful dark gray color as the living black heron has”. Alexander Wetmore (in Ackermann 1964: 62), when asked about the possible identity of the feathers in communication by letter, could only indicate that, due to the wide and loose barbs, it could be tail coverts. He also suggested Ackermann should include a mention of the feathers in a publication about his finds due to its scientific interest, on which Ackermann commented that “although the species of the bird to which it belonged cannot be determined, it deserves to be highlighted due to its rarity and preservation”.
Although Ackermann mentioned a single feather in the text, two photographs in black and white and with scale were included. They clearly show two distinct specimens, and an excerpt from Wetmore’s letter also mentions them in the plural. The specimens measure approximately 6 cm and 7 cm. Prado et al. ( 2016a ) considered them possible semiplumes or contour feathers. The whereabouts of these specimens are unknown.
Fossils of cnidarians, bivalves, gastropods, cartilaginous fishes, and several plant species were found in the same locality, which indicates an estuarine environment (Távora et al. 2010 , Góes et al. 1990 ).
Feathers from the Crato and Tremembé Formations: GP/2E-9484 (A); GP/2E-9485 (B); GP/2E-9486 (C); GP/2E-9494 (D); LEG0766 (E); LEG0896 (F); LEG0885 (G); LEG0886 (H); MHNCE/FOS-001 (I); DGM 1-A (J–K); GP/2E-8125 (L); GP/2E-8126 (M); GP/2E-8127 (N) (A–D, M–N: R.S. Nascimento; E–H, L: G.M.E.M. Prado; I: M.A. Crozariol; J–K: R.C. Silva). Scale bar: 5 mm, except E–H.
2.4.5. Eggs
The paleoological record of birds in Brazil includes just two reports, representing both Mesozoic and Cenozoic sediments. No ootaxa were erected. Reports of eggshells in the archeological literature (e.g., Simonsen et al. 1983 –1984, Prous 2009 , Jacobus and Rosa 2013 ) generally do not specify whether they are from reptiles or birds (though the latter is more probable). Records with attributions so far belong to paleognaths. Unspecified avian mentions include that of Prous et al. ( 1996 –1997) from Lapa do Dragão in Montalvânia, Minas Gerais, and the two indeterminate egg types reported by Mentz Ribeiro ( 1983 ) from the Anápio de Oliveira (RS-RP-83) site in Vera Cruz, Rio Grande do Sul.
2.4.5.1. Mesozoic eggs
192. †Ornithothoraces indet. (LPRP-USP 0359)
Late Cretaceous – SP
“ovo fossilizado afim a Aves” – Marsola et al. 2012: 49.
“ovo de aves Enantiornithes” – Marsola 2013: 25, 49–55, figs 10, 23–27.
Ornithothoraces – Marsola et al. 2014: 563–566, 2.
“ave” – Reis et al. 2014a: 184.
“avian egg is similar to those of the genus Enantiornithes ” [sic] – Reis et al. 2014b: 794.
Ornithothoraces – Martinelli and Teixeira 2015: 147, 149.
Ornithothoraces – Langer et al. 2022: 19–20.
Marsola et al. ( 2014 ) reported an egg attributed to the Ornithothoraces from the Late Cretaceous of Adamantina [Vale do Rio do Peixe] Formation, being the first recorded occurrence of a Mesozoic fossil egg for Brazil. The fossil was previously provisionally described by Marsola et al. ( 2012 ) and Marsola ( 2013 ). It was found in July 2011 in a site along the SP-270 road near Álvares Machado.
A combination of Scanning Electron Microscopy, Wave Dispersion Energy analyses, and Computed Tomography was used to determine its taxonomic identity and structural features of its biomineralized tissues. The specimen (LPRP-USP 0359) is a nearly complete, slightly compressed egg, with its main axes measuring 31.4 mm x 19.5 mm. It is one of the smallest known Mesozoic avian eggs. The deformation was probably caused by lithostatic compression during sedimentation. There is a small polar portion damaged, possibly by erosion during exposure in the outcrop. The 125.5-μm-thick shell is externally smooth with rounded pore openings and incorporates three structural layers of similar thickness with both prismatic and aprismatic boundaries. It is somewhat fragile in comparison to the eggs of modern neognaths and paleognaths and represents one of the thinnest shelled Mesozoic avian eggs known. There are no signs of predation, hatching, or trampling, nor embryonic remains (Marsola 2013 ).
The egg is very similar to those of enantiornitheans from the Late Cretaceous of Bajo de la Carpa Formation (Río Colorado Subgroup) of Argentina. Marsola et al. ( 2012 ) and Marsola ( 2013 ) initially attributed it to Enantiornithes, while Marsola et al. ( 2014 ) preferred an affinity with basal Ornithothoraces. Furthermore, the similarity of both depositional contexts suggests a shared preference regarding breeding and nesting environments. The paleoenvironment of the Vale do Rio do Peixe Formation is very similar to that of the Bajo de la Carpa Formation, a semi-arid landscape with eolian dunes interspersed with wide, shallow, and slightly anastomosing ephemeral river systems that were subject to seasonal fluctuation.
2.4.5.2. Cenozoic eggs
193. †Aves indet. (MN 4705-V)
Azevedo and Carvalho ( 1998 ) reported several small and thin eggshell fragments (MN 4705-V) from the upper Tremembé Formation. Analysis through Scanning Electron Microscopy and microscopes of the fractured and polished surfaces of mainly radial sections showed a structural arrangement that strongly resembles the typical ratite pattern. Four layers could be differentiated, an innermost zone of radial calcite plates (~0.03 mm), a basal tabular crystallite aggregates layer (~0.14 mm), a spongy layer (~0.19 mm), and a thin external zone (~0.02 mm), with pores distributed randomly between the shell units.
Previously, Garcia ( 1993 ) reported fragments of eggshells from the Taubaté Basin in the UNG collection but did not attribute them to any specific group of vertebrates, and, so far, the study of Azevedo and Carvalho constitutes the only secure record of avian eggs from this unit.
194. Rhea americana (Linnaeus) (Eggs)
Mentz Ribeiro ( 1972 ) reported eggshells from the Schneider (or Bom Jardim Velho) (RS-C-14) rock shelter in São Sebastião do Caí, Rio Grande do Sul. He noted this is the first record for the archeology of that State.
“cascas de ovos” [?; in part?] – Schorr 1976: 97.
“casca de ovos da mesma ave [ Rhea americana ]” [in part?] – Schmitz 1980: 200.
“cáscaras de huevo de esta ave [ Rhea americana ]” – Schmitz et al. 1981: 93.
“Cascas de ovos” [?; in part?] – Jacobus and Schmitz 1983a: 58, 8L–P.
“Casca de ovos” [?; in part?] – Jacobus and Schmitz 1983b: 268, 7L–P.
“casca de ovos” [?; in part?] – Jacobus (in Schmitz and Machado 1987 ): 167.
“casca de ovo de Rhea americana ” – Schmitz et al. 1989: 154.
“cascas de ovos da mesma ave [ Rhea americana ]” [in part?] – Barbosa et al. 1990: 89.
“ovos das emas” [in part?] – Schmitz 1990: 110.
“ema, de cujos ovos apareceram abundantes pequenos fragmentos de casca” – Schmitz 1999: 93.
Rhea americana – Rosa 2004: 260.
“ovos de ema” – Pacheco 2008: 25.
“ovos de emas” – Guimarães 2011: 109.
Schorr ( 1976 ) reported eggshells among the remains from the GO-JA-01, GO-JA-14, and GO-JA-20 rock shelters in Serranópolis, Goiás, without determining its species of origin, as did Jacobus and Schmitz ( 1983a , 1983b ) from GO-JA-01. Later mentions in the literature of eggshells from these sites generally attribute them to Rhea americana and note they are a common occurrence (Schmitz 1980 , 1999 , Schmitz et al. 1989 for GO-JA-01, Rosa 2004 ), in contrast with the scarce osteological remains. These materials are deposited in the IAP/Unisinos collection.
Junqueira and Malta ( 1981 –1982) reported human-made adornments made of rhea eggs from the late Holocene of Lapa da Hora in Januária, Minas Gerais.
“cascas de ovo de ema” – Barbosa et al. 1982: 75.
Barbosa et al. ( 1982 ) reported eggshells from the late Holocene of the GO-RS-01 rock shelter in Monte do Carmo, Goiás.
“Casca de ovo de ema: ( Rhea americana )” [in part] – Mentz Ribeiro 1983: 20.
“Casca de ovo de ema” [in part] – Mentz Ribeiro 1983: 22.
“Casca de ovo de ema” [in part] – Mentz Ribeiro 1983: 26.
“casca de ovo de ema ( Rhea americana )” [in part] – Mentz Ribeiro 1991b: 79.
“casca de ovo de ema ( Rhea americana )” [in part] – Mentz Ribeiro 1991b: 119.
“ovos de ema” – Mentz Ribeiro 1995: 12.
Mentz Ribeiro ( 1983 ) reported small eggshell fragments from the latest Holocene of Anápio de Oliveira (RS-RP-83) site in Vera Cruz, Rio Grande do Sul. The material consists of two fragments from layer IV (oldest), seven from layer V, and 21 from layer VI.
“Ovos de Rhea americana (fragmentos)” – Mentz Ribeiro et al. 1989: 75.
“casca de ovo de ema” – Mentz Ribeiro 1991a: 114.
“Fragmentos de ovos de Rhea americana ” – Mentz Ribeiro and Ribeiro 1999: 29.
“eggs of Rhea americana ” [in part?] – Prous and Fogaça 1999: 27.
“fragmentos de casca de ovos de Rhea americana ” – Rosa 2006f: 2.
Rhea americana – Rosa 2007a: 40.
Rhea americana – Rosa 2009: 152.
Rhea americana [in part] – Dias 2012: 15.
Rhea americana – Hadler et al. 2013: 121.
“cascas de ovos” [in part] – Jacobus and Rosa 2013: 250.
Mentz Ribeiro et al. ( 1989 ) reported eggshell fragments from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Rosa ( 2009 ) reported this material to consist of 5,165 eggshell fragments (1,675 from square A6, 364 from A7, 379 from B5, 529 from B6, 621 from B7, 392 from C5, 13 from C6, 364 from C7, 160 from D5, 507 from D6, and 161 from D7). Remains from square A6 were detected in the site’s three occupation periods. The material is deposited in the CEPA collection.
“fragmentos de ovo de ema” – Vilhena Vialou and Vialou 1989: 39.
Vilhena Vialou and Vialou ( 1989 ) reported eggshells from the Santa Elina rock shelter in Jangada, Mato Grosso.
“cascas de ovos de ema, Rhea americana ” – Silva 2003: 291.
Silva ( 2003 ) reported eggshells from layers I (a fragment of historical age) and III (four fragments) of the Ilha de Sorobabel site in Itacuruba, Pernambuco.
“fragmentos de ovos de ema ( Rhea americana )” – Brentano et al. 2006: 215.
Brentano et al. ( 2006 ) reported eggshells from the RS-LC-97 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
“[Da ema] fragmentos de ovos” – Rosa 2006b: 228.
Rosa ( 2006b ) reported eggshells from the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. Bones were also found. The material is deposited in the IAP/Unisinos collection.
“ovos de ema” – Prous 2007: 35.
Prous ( 2007 ) mentioned rhea eggs among the archeofaunal remains of Dalpiaz, Batinga, and Ivoti rock shelters in Southern Brazil.
“fragmentos de casca de ovo de ema (Rheidae)” – Farias et al. 2016: 4.
Farias et al. ( 2016 ) reported eggshell fragments from the RS-TQ-140 site in Tabaí, Rio Grande do Sul, dated 3060±30 CAL years BP. The site was excavated in December 2013 by the staff of Grupo de Pesquisa em Educação Patrimonial e Arqueologia (GRUPEP) of Universidade do Sul de Santa Catarina.
195. cf. Tinamidae (Eggs)
Rosa ( 2009 ) reported 415 possible tinamid eggshell fragments (74 from square A7, 41 from B6, 246 from B7, 24 from C5, 15 from C7, and 15 from D5) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Other 159 indeterminate avian eggshell fragments were also found. The material is deposited in the CEPA collection.
2.4.6. Osteological remains and mummifications
Osteological remains make up the bulk of the known prehistoric bird material for Brazil. Most extinct taxa were described from truly fossilized remains, but many subfossils are also known, especially from archeological sites. Mummified remains associated with these sub-recent materials are rarer and only mentioned in the literature on a few occasions.
Aves Linnaeus
196. †Aves indet. 1 (SMF)
Early Cretaceous – CE
“partial ‘wing’” – Naish et al. 2007: 528, pl. 25D.
cf. Aves [in part] – Bittencourt and Langer 2011: 28–29, 38.
“Three other asymmetrical feathers found in association with bone fragments” – Sayão et al. 2011: 198.
“possíveis aves” [in part] – Leite and Hessel 2011: 2.
“materiais osteológicos de aves” [in part] – Pinheiro and Fernandes-Ferreira 2014: 125.
“avian skeletons associated with poorly preserved feathers” [in part] – Carvalho et al. 2015a: 2.
“presumed carpal bones associated with three asymmetrical feathers” – Carvalho et al. 2015b: 162.
“materiais isolados de aves” [in part] – Metello 2017: 6.
“enantiornithines” [?; in part] – Pittman et al. 2020: 61.
Naish et al. ( 2007 ) briefly described and depicted an unaccessioned specimen from the Crato Formation in the SMF collection. It comprises a single large asymmetric “primary” flight feather similar to that described by Martins Neto and Kellner ( 1988 ), five smaller asymmetric feathers, and two unidentifiable, presumed carpal bones [20] . The longest and smallest feathers are 81 mm and 14 mm long, respectively.
197. †Aves indet. 2 (UFRJ-DG 06 Av)
Late Cretaceous – SP
Aves indet. – Azevedo et al. 2007: 546, 2R–S.
cf. Aves – Bittencourt and Langer 2011: 38.
Aves indet. – Candeiro et al. 2012: 618.
Aves cf [?; in part?] – Brum and Simbras 2014: 7.
Aves cf [?; in part?] – Brum and Simbras 2015: 119.
Aves indet. – Martinelli and Teixeira 2015: 147.
Aves indet. – Alves et al. 2016: 87–88, 4I–J.
Azevedo et al. ( 2007 ) reported a bone fragment (UFRJ-DG 06 Av) about 1 mm long from the Jales locality of Adamantina [Vale do Rio do Peixe] Formation in Alfredo Marcondes, São Paulo. They tentatively attributed it to the distal end of a phalanx of an indeterminate bird. The fossil was also described and depicted by Alves et al. ( 2016 ).
198. †Aves indet. 3 (CPPLIP 470)
Candeiro et al. ( 2012 ) assigned to an indeterminate Aves a complete, robust isolated pedal phalanx 1 of left digit II [21] (CPPLIP 470) from the Ponto 1 do Price site of Peirópolis locality (Marília Formation) in Uberaba, Minas Gerais. It was discovered in 2009 and briefly mentioned by Candeiro et al. ( 2010 ). The authors noted similarities with the larger specimen MACN PV RN 1107 described by Agnolín and Martinelli ( 2009 ) from the Late Cretaceous Los Alamitos Formation of Río Negro Province, Argentina, including features reminiscent of derived neornithine predatorial birds (such as falconiforms). However, Candeiro et al. ( 2010 ) tentatively assigned it to Neornithes indet. and Candeiro et al. (2012) to Aves indet. only.
199. †Aves indet. 4 (CPPLIP 481)
Candeiro et al. ( 2012 ) assigned a small, isolated bone (CPPLIP 481) from the Ponto 1 do Price to an indeterminate Aves and identified it as a possible pedal ungual phalanx lacking the proximal end. Its position on the foot could not be determined due to its incomplete nature.
†Enantiornithes Walker
200. †?Enantiornithes indet. (MURJ)
Early Cretaceous – CE
Figs 15–16
“probable enantiornithine bird” – Naish et al. 2007: 528–529, pl. 26.
“a nearly complete and excellently well preserved skeleton of a bird” [?] – Herzog et al. 2008: 38.
“bird, possibly enantiornithine” – Close et al. 2009: 617.
“enantiornithine bird” – O’Connor and Dyke 2010: 19.
Enantiornithes indet. [in part] – Bittencourt and Langer 2011: 28–29, 38.
“ave enantiornitínea” – Campos 2011: 90.
“supposed presence of an enantiornithine bird” – Sayão et al. 2011: 203.
“possivelmente aves euenantiornitiformes” – Leite and Hessel 2011: 2.
Enantiornithes indet. – Bittencourt and Langer 2012: 305–306.
“possível enantiornitíneo” – Pinheiro and Fernandes-Ferreira 2014: 125.
“avian skeletons associated with poorly preserved feathers” [in part] – Carvalho et al. 2015a: 2.
“poorly preserved articulated specimen” – Carvalho et al. 2015b: 162.
“materiais isolados de aves” [in part] – Metello 2017: 6, 5.
“enantiornithines” [in part?] – Pittman et al. 2020: 61.
Naish et al. ( 2007 ) preliminary described a partial skeleton associated with feathers of a probable enantiornithean from the Crato Formation held in the MURJ private collection in Japan. The poorly preserved fossil is somewhat crushed and partially articulated, preserved in part and counterpart. Many of the bones are represented only by external molds. The skull is preserved in palatal view and is incomplete, with a braincase, mid-line palatal elements, and possible rod-like jugal bar discernible in the photographs available to the authors. About 25 vertebrae are preserved, including cervical (several in ventral view), dorsal, and caudal. The dorsal vertebrae show features comparable to that of euenantiornitheans. At least three caudal vertebrae are preserved, two of which are articulated. Fragments of other caudal vertebrae are also present. A presumable pygostyle was not preserved. Part of the ilium and a possible ischium were tentatively identified. A hindlimb, possibly the left one, is the best-preserved part of the specimen, comprising the femur, tibiotarsus, tarsometatarsus, and hallux. The last appears proportionally large, fully reversed, and retaining its ungual claw sheath. Feather impressions are present alongside the tarsometatarsus and adjacent to the femur. The specimen is 156 mm long from the rostralmost preserved tip of the skull to the last preserved caudal vertebrae, about the same size as the fringillid Fringilla coelebs .
Herzog et al. ( 2008 ) mentioned a nearly complete and excellently well-preserved skeleton rumored to have been sold and smuggled out of Brazil, with unknown whereabouts. It is unclear whether this refers to the MURJ specimen, though this seems likely.
201. †cf. Enantiornithes (CPPLIP 482)
Late Cretaceous – MG
cf. Enantiornithes gen. et sp. indet. – Candeiro et al. 2012: 620, 1C.
Enantiornithes indet. – Candeiro et al. 2013: 17, 18, 19.
Enantiornithes – Candeiro 2015: 165, 36J.
cf. Enantiornithes gen. et sp. indet. – Martinelli and Teixeira 2015: 147, 10C.
Theropoda indet. [?; in part] – Pereira et al. 2015: 175.
From the Ponto 1 do Price site, Candeiro et al. ( 2012 ) assigned an incomplete left metatarsal III lacking its proximal portion (CPPLIP 482) to a possible enantiornithean.
202. †Enantiornithes indet. (spp.) (William’s Quarry)
Late Cretaceous – SP
Enantiornithes – Alvarenga and Nava 2005: 20.
“at least three indeterminated different Enantiornithes” – Agnolín and Martinelli 2009: 42.
Enantiornithes – Alvarenga and Höfling 2011: 119.
Enantiornithes indet. – Bittencourt and Langer 2011: 33, 38.
Enantiornithes indet. – Bittencourt and Langer 2012: 310–311.
Enantiornithes indet. – Candeiro et al. 2012: 618.
Enantiornithes – Marsola et al. 2012: 49.
“pequenas aves associadas aos Enantiornithes” [in part?] – Nava 2012: 47.
“pequenas aves relacionadas aos Enantiornithes” [in part?] – Nava 2013: 253.
Aves cf [?; in part?] – Brum and Simbras 2014: 7.
Aves cf [in part?] – Brum and Simbras 2015: 119.
“pequenas aves relacionadas aos Enantiornithes” [in part?] – Nava 2015: 138.
“indeterminate enantiornithine birds” – Carvalho et al. 2015b: 161.
Enantiornithes indet. – Martinelli and Teixeira 2015: 147.
“enantiornithine birds” [in part] – Nava et al. 2015: 73.
Enantiornithes [in part] – Alves et al. 2016: 87–88.
“enantiornithine birds” [in part] – Chiappe et al. 2018a: 170.
“enantiornithine birds” [in part] – Chiappe et al. 2018b: 105.
“enantiornithine birds” [in part] – Chiappe et al. 2019: 232–233.
“enantiornithines” [in part] – Pittman et al. 2020: 61.
Enantiornithes [in part] – Langer et al. 2022: 19, 9.
Enantiornithes [in part] – Paixão et al. 2023: 74.
Alvarenga and Nava ( 2005 ) reported enantiornithean remains from the William’s Quarry site (Chiappe et al. 2018a ) of Adamantina Formation in Presidente Prudente, São Paulo. They constitute the first avian bones of the Mesozoic to be reported for the country. The first material was collected two months before their public announcement on 10 August 2005. It is represented by several dozens of three-dimensionally preserved and delicate bones, isolated and partially articulated, of about three or four different taxa morphologically very close to Enantiornis leali[22] from the Late Cretaceous of El Brete, Argentina, but much smaller, of size comparable to small passeriforms. Differences in phalanx thickness indicate that disparate taxa are represented (Alvarenga in Castilhos 2005 ). Additionally, one form has four and the other three toes (Lopes 2005 ). Despite their great diagnostic value, the bones were difficult to associate, which prevented the description of new taxa.
Alvarenga and Höfling ( 2011 ) described the still-under-study material as including several bones, many as part of incomplete skeletons that represent at least four genera and species that did not seem to have teeth and were hummingbird or sparrow-sized.
The material still awaits formal description, but several bones were depicted in contemporaneous news media (e.g., Castilhos 2005 , Lopes 2005 ), including a humerus with associated radius and ulna and other fragments (MHNT-VT-5240), a complete left coracoid (MHNT-VT-5241), and an incomplete tarsometatarsus (MHNT-VT-5242). In addition, toothless skulls, vertebrae, hindlimb elements, and other fragments were also mentioned by Lopes ( 2005 ).
“pequenas aves associadas aos Enantiornithes” [in part?] – Nava 2012: 47.
“pequenas aves relacionadas aos Enantiornithes” [in part?] – Nava 2013: 253.
Aves cf [?; in part?] – Brum and Simbras 2014: 7.
Aves cf [in part?] – Brum and Simbras 2015: 119.
“pequenas aves relacionadas aos Enantiornithes” [in part?] – Nava 2015: 138.
“enantiornithine birds” [in part] – Nava et al. 2015: 73.
Enantiornithes [in part] – Alves et al. 2016: 87–88.
“enantiornithine birds” [in part] – Chiappe et al. 2018a: 170.
“enantiornithine birds” [in part] – Chiappe et al. 2018b: 105.
“enantiornithine birds” [in part] – Chiappe et al. 2019: 232–233.
“enantiornithine birds” [in part] – Wu et al. 2019: 221.
“enantiornithines” [in part] – Pittman et al. 2020: 61.
“enantiornithines” [in part] – Wu et al. 2021: 1–12, figs 1–4.
“enantiornithines” [in part] – Chiappe et al. 2022: 1–12, 1.
Enantiornithes [in part] – Langer et al. 2022: 19, 9.
Enantiornithes [in part] – Paixão et al. 2023: 74.
More material from William’s Quarry was excavated in the subsequent years after the publication of Alvarenga and Nava ( 2005 ). Nearly 1,000 articulated and isolated bird bones were discovered, of which the diagnosable ones belong to Enantiornithes (Wu et al. 2019 , Chiappe et al. 2022 ). The discovered material includes cranial, such as isolated rostra, mandibles, and crania, and postcranial elements, including associated sacral and lumbar vertebrae forming a synsacrum, furcula, and partially fused carpometacarpus and tarsometatarsus, representing at least three small to medium-sized taxa (Nava 2012 , 2013 , 2015 , Nava et al. 2015 , Chiappe et al. 2018a , 2018b , 2019 ). This material is deposited in the MPM collection and is under preparation at that institution and LACM (Fonseca 2019b , SAPE 2020 ).
While the taxonomic description of the material is still pending, other morphological and developmental studies were published. Nava et al. ( 2015 ) briefly described two three-dimensionally preserved, dorsally V-shaped toothed premaxillae, which were found associated with postcranial elements of unquestionably enantiornithean identity.
Wu et al. ( 2021 ) μCT scanned two sets of premaxillae (MPM-90 and MPM-373) and the rostral portion of a left dentary (MPM-351) exquisitely preserved in three dimensions to better understand the occurrence of polyphyodonty in Mesozoic birds. The two sets of premaxillae are very similar, and likely represent the same species or very closely related taxa, despite MPM-373 being significantly smaller and slightly more compressed laterally. Both have four tooth positions with most of their conical and slightly recurved teeth preserved in place, with the first two teeth of a smaller size. This differs from the material described by Nava et al. ( 2015 ), in which the first pair of teeth is much larger than the last. The dentary preserves the first four teeth (the fourth being somewhat broken), which are also conical and slightly recurved, with a similar size and evenly spaced. The results showed preservation of replacement teeth and their tooth families, and a three-dimensional reconstruction of the replacement tooth rows showed a conserved alternating pattern, as observed in other archosaurs. Previously, Wu et al. ( 2019 ) briefly reported the results on specimens MPM-90 and MPM-351.
Chiappe et al. ( 2022 ) studied a nearly undistorted basicranium (MPM-334-1) that substantially expanded the knowledge of the early evolution of avian braincase, brain, and inner ear. It presents traits regarded as exclusive to Neornithes and suggests they may have originated before the split between the enantiornitheans and crown birds over 140 million years BP. The fossil appears to be skeletally mature and is small, falling within the size range of the smallest known enantiornithines, as well as some extant hummingbirds and small passerines.
†Euenantiornithes Chiappe
† Cratoavis Carvalho, Novas, Agnolín, Isasi, Freitas & Andrade
203. † Cratoavis cearensis Carvalho, Novas, Agnolín, Isasi, Freitas & Andrade
Early Cretaceous – CE
Figs 17A–B, 30A
Type locality: Pedra Branca Mine, Nova Olinda County, Ceará State, Brazil (7° 6’51.9” S and 39° 41’46.9” W).
Etymology: Cratoavis , with the Latin avis , “Crato bird”, and cearensis , referring to the state of Ceará.
Indeterminate genus and species – Carvalho et al. 2015a: 1–5, supl. 1–19, figs 1–2, supl. figs 1–7.
Cratoavis cearensis nov. gen. et sp. [new genus and species] – Carvalho et al. 2015b: 161–171, figs 3–4.
Cratoavis cearensis – Metello 2017: 7–8, 28, 30, figs 1, 16.
Cratoavis cearensis – Agnolín et al. 2017b: 2, 1A.
“UFRJ-DG 031Av” – Knoll et al. 2018: 6–7.
“ Cratoavis ” – O’Connor 2020: 157.
Cratoavis cearensis – Foth 2020: 176–177.
“UFRJ-DG 031Av” – O’Connor et al. 2020: 37, 39, 40.
“ Cratoavis (valid?)” – Pittman et al. 2020: 61.
Carvalho et al. ( 2015a ) described an articulated skeleton associated with feathers of a minute euenantiornithean from Mina Pedra Branca in Nova Olinda, Ceará. It was found by Cleuduardo Laurentino Dias, Devânio Ferreira Lima, and Antonio Josieudo Pereira Lima. Later in the same year, they described it further and erected the new genus and species Cratoavis cearensis for the material (Carvalho et al. 2015b ).
The specimen (UFRJ-DG 031 Av, holotype) is a nearly complete skeleton preserved in part and counterpart, about 6 cm long from the snout to the tip of the pygostyle. The skeleton is exposed in lateral view, except for the proximal caudal vertebrae and pygostyle (including the rectrices), which are exposed in dorsal view. The skull and neck are exposed on the left side, rotated ventrally regarding the rest of the body, whereas most of the remaining skeleton elements are exposed on the right side. The bones have a dark-brown coloration and are preserved in three dimensions, but as commonly in fossils preserved in part and counterpart, several elements are cracked, with crushing and displacement in some of them (e.g., in the skull and jaw). The smaller counterpart shows better preservation but contains only imprints of the skull, forelimbs, and pectoral girdle fragments.
Cratoavis cearensis (UFRJ-DG 031 Av): skeleton in counterslab (A) and partial skeleton in slab (B); Kaririavis mater (UFRJ-DG 116 Av): partial foot with associated feathers in slab (C) (I.S. Carvalho). Scale bar: 5 mm.
The identified elements, both as bones or impressions of varying degrees of preservation, include parts of the poorly-preserved skull (parietal, frontal, nasal, and lacrimal bones), sclerotic ring, maxilla with minute alveoli (which indicates it was toothed), vertebrae (five cervical preserved in articulation, six dorsal, and eight free caudal, as well as sacral), pygostyle (composed of eight fused vertebrae), most of the right dorsal ribs and the distal end of some left ones, sternal ribs, both ilea, pubes, both coracoids, left scapula, both humeri, both radii, both ulnae, both carpometacarpi, left-wing phalanges (represented by the alular and major digits, including unguals, and only the base of the first phalanx of the minor digit), both femora, both tibiotarsi, both metatarsals (best represented by right I, II, III, and IV ones), and pedal phalanges of both feet (represented by digits I, II, III, and IV, including unguals). In addition, long, thin bones in the abdominal region are probably part of a gastralium. The ischia and the sternum were not preserved. Knoll et al. ( 2018 ) noted that the latter was presumably completely cartilaginous.
Different elements of the plumage were preserved, most as impressions (Metello 2017 ). The most remarkable are the paired, dorsoventrally depressed, rachis-dominated, 79.9-mm-long symmetrical rectrices, roughly 30% longer than the length of the skeleton, similar to that observed among other enantiornitheans and confuciusornithiforms. Their proximal portion was interpreted as the calamus, inserted on the third proximal pygostyle vertebra. The preservation in relief helped recognize key characters in this feather morphotype: a narrow groove extending from the base to the distal tip; the robust scapus nearly flat at the calamus, becoming slightly convex in the rachis at mid-length (with its reconstruction indicating it was nearly 8-shaped in cross-section), flattening again distally where it becomes vaned; the first recognizable barbs of the pennaceous vanes appearing at nearly mid-length of the scapus, increasing in size distally; and no visible signs of interlocking barbules. According to O’Connor ( 2020 ), the three-dimensional specimens preserved in Cretaceous Burmese amber (showing that the wide rachis observed in compression fossils was originally C-shaped with a thickness of 3–10 μm) indicate that rectrices of C. cearensis are dorsally exposed and flattened. However, Foth ( 2020 ) noted that this particular morphology could be the result of miniaturization and that Carvalho et al. provided no compelling evidence that the rectrices were de facto preserved in dorsal view (other than the caudal vertebrae and pygostyle being preserved in the same view). The longitudinal groove most likely represents the ventral furrow of the rachis and not a dorsal groove, in analogy to modern pennaceous feathers. The third proximal part of the rachis bears a row of five transversal lunar dark granulate bands, interpreted as remnants of an ornamental color pattern, preserved as carbonized traces (Metello 2017 ). The presence of this kind of pattern reinforces the interpretation that such elongated rectrices could have been sexually dimorphic and associated with sexual display, specific recognition, or visual communication among early birds. Small brownish contour feathers with pennaceous and plumulaceous vanes cover the wings and body, including a crown at the top of the head. Prado ( 2017 ) speculated they might consist of melanosomes and were possibly grayish or reddish-colored. Ten asymmetrical secondary remiges were incompletely preserved (longest right one 10.3 mm, longest left one 8.6 mm), and asymmetrical alular feathers are also observable, with at least three preserved on the left hand. Feathers on the hind limbs were not observed. There are probable soft tissues preserved around the skull, in some cases appearing like a dotted surface similar to skin fragments, and small plumes appearing like a brownish tinge. Elongate, yellowish-white muscle fibers are preserved bordering the femur.
The ontogenetic state of the specimen is uncertain. Carvalho et al. ( 2015b ) pointed out the very small body size and unfused tarsometatarsus, tibiotarsus, and carpometacarpus as evidence in favor of it being a young individual. However, the fused vertebral centra, well-defined epiphyses of long bones, extensive pubic apron, and absence of surface pitting indicate it was probably an adult individual. A well-developed plumage is also contrasting, especially the long rectrices, similar to those observed in adult modern birds. The early ontogenetic emergence of these tail feathers is also indicated by a well-developed plumage, especially the ornamental rectrices, in young enantiornitheans from the Jehol Group (Early Cretaceous of northeast China). If correct, this could indicate that significant differences in plumage development probably existed between enantiornitheans and modern birds. It was referred to as an immature specimen by Knoll et al. ( 2018 ), O’Connor ( 2020 ), and O’Connor et al. ( 2020 ).
Carvalho et al. ( 2015a , 2015b ) classified it as a euenantiornithean of unresolved phylogenetic position and highlighted similarities with the genera Pengornis (Early Cretaceous of Jiufotang Formation, China) and Eoenantiornis (Early Cretaceous of Yixian Formation, China). Agnolín et al. ( 2017b ) referred to it as a basal enantiornithean. O’Connor ( 2020 ) referred to the taxon as “ Cratoavis ” and Pittman et al. ( 2020 ) as “ Cratoavis (valid?)”, but neither of them developed it further. The phylogenetic analysis of Wang et al. ( 2022 ) recovered it as a sister taxon of the genus Gretcheniao (Early Cretaceous of Yixian Formation, China).
Ornithuromorpha Chiappe
† Kaririavis Carvalho, Agnolín, Rozadilla, Novas, Ferreira Gomes Andrade & Xavier-Neto
204. † Kaririavis mater Carvalho, Agnolín, Rozadilla, Novas, Ferreira Gomes Andrade & Xavier-Neto
Early Cretaceous – CE
Figs 17C, 30B
Type locality: Pedra Branca Mine, Nova Olinda County, Ceará State, Brazil (7°7’25”S, 39°42’7”W).
Etymology: Kaririavis , with the Latin avis , “Kariri bird”, referring to the Karirí people of northeastern Brazil, and mater , Latin for “mother”, as it is the oldest ornithuromorph known for South America.
“early Cretaceous birds from Brazil” – SAPE 2020: 3.
“modern birds from Cretaceous beds from Brazil and Argentina” [in part] – SAPE 2021: 3.
Kaririavis mater gen. et sp. nov. [new genus and species] – Carvalho et al. 2021: 1–12, figs 2–4, 6.
Carvalho et al. ( 2021 ) described a small, partial isolated foot from the Crato Formation of Mina Pedra Branca in Nova Olinda, Ceará, as the new genus and species Kaririavis mater . A study of the fossil had already been reported in the previous year (SAPE 2020 ) but with no details other than its age and country of origin.
The fossil (UFRJ-DG 116 Av) is preserved as part and counterpart and includes partial right foot, a nearly complete tarsometatarsus (exposed in plantar view), and several apparently unconnected pedal phalanges, tentatively identified as digit II phalanges 1 and 2, digit III phalanges 1 to 3, two indeterminate phalanges of digit IV, and ungual phalanx of digit II covered with a keratinous sheet. At least 10 poorly preserved dark-brownish contour feathers were also preserved in close association, which the authors believed probably belonged to this individual.
The sparrow-sized bird was referred to Ornithuromorpha. Its unique foot morphology precludes a clear knowledge of its habits and the paleoecological niche it occupied, but the morphology of its tarsometatarsus strongly suggests it may belong to an unknown ornithuromorph clade with terrestrial habits, in contrast with most Early Cretaceous ornithuromorphs, which are interpreted as semiaquatic.
Up to its description, the knowledge of Early Cretaceous ornithuromorphs was almost restricted to Asia and absent from Gondwanan landmasses. The discovery of K. mater considerably expands the paleobiogeographic distribution of Ornithuromorpha. It is one of the oldest worldwide occurrences, as well as the first from Gondwana, with similar age to species from China. The record reinforces the idea that the diverse early Aptian basal ornithuromorphs known as “Jehol Birds” were probably not unique to eastern Asia but represent an example of biota from multiple lagerstätte sites serendipitously first found in that continent. Despite Early Cretaceous birds still being insufficiently known, K. mater exhibits a very different morphology when compared with contemporary Asian avifaunas, which is possibly a result of biogeographic differences and indicates that the phylogenetic diversity of Mesozoic avifaunas of Gondwana is likely far greater than what has been discovered so far.
Neornithes Gadow
205. †Neornithes indet. 1
Taranto and Bergqvist ( 2009 ), Taranto et al. ( 2009a ), and Taranto and Bergqvist ( 2010 ) associated with Ralliformes (=Cariamiformes?) the distal end of a right femur (UFRJ 01-AV) from the São José de Itaboraí. Taranto and Bergqvist ( 2010 ) described the bone as having a similar size to that of a pigeon, a comparison associated with a tibiotarsus (UFRJ 02-AV) in the two previous publications (see below). Both fragments were long stored in the Departamento de Geologia of UFRJ.
206. †Neornithes indet. 2
Early Eocene – RJ
“muito similar ao Paleopsilopteris itaboraiensis ” – Taranto and Bergqvist 2009: 51R.
“nova forma de Phorusrhacidae” – Taranto et al. 2009a: 287.
“tamanho e características muito próximas a tauracos (Musophagiformes) e urias (Charadriiformes), mas não foi associado a estes grupos pela ausência da porção proximal, onde está a maior parte das sinapomorfias” – Taranto and Bergqvist 2010: 118.
Taranto and Bergqvist ( 2009 ) and Taranto et al. ( 2009a ) reported the distal end of a right tibiotarsus (UFRJ 02-AV) from the São José de Itaboraí Basin. They noted that the bone is similar to that of Paleopsilopterus itaboraiensis , but better preserved and close in size to that of a pigeon. It belongs to an adult individual, suggesting another related form in the basin. However, Taranto and Bergqvist ( 2010 ) associated the “size of a pigeon” with a femur (see above) and described the tibiotarsus as similar in size and form to Musophagiformes and Charadriiformes but did not associate it to any of these groups due to the lack of the bone’s proximal end, where most synapomorphies are concentrated.
207. †Neornithes indet. 3
Mayr et al. ( 2011a ) reported a right carpometacarpus (MN 4115-V) from the São José de Itaboraí Basin. Due to the assignment of two humeri and a coracoid to Itaboravis elaphrocnemoides (see Cariamiformes), they pointed out that this species seems to be among the most abundant small birds in Itaboraí, and this could support the attribution of this carpometacarpus of comparable size to it. However, the Itaboraí material also includes tibiotarsi of four small to medium-sized taxa (see below), which demonstrates a considerable diversity and, unlike one of the humeri and the coracoid, which were found in the same matrix block, there is no evidence beyond the size to support this attribution to I. elaphrocnemoides . It has a distinctive morphology, unlike any other known avian taxon, and most closely resembles tinamids. However, it differs from them in several other characteristics, and clearly differs from the European Paleogene genus Elaphrocnemus .
208. †Neornithes indet. 4
Mayr et al. ( 2011a ) reported the distal end of a left tibiotarsus (MN 4119-V) from the São José de Itaboraí Basin. It belongs to a small bird, much smaller than Itaboravis elaphrocnemoides , perhaps Eutreptodactylus itaboraiensis , the only bird of comparable size known from the same locality and whose holotype came from the same set of bones. However, because the holotype of E. itaboraiensis is a tarsometatarsus and has been lost, further comparisons are not possible for a reliable attribution of MN 4119-V.
209. †Neornithes indet. 5
Mayr et al. ( 2011a ) reported the distal end of a right tibiotarsus (MN 4116-V) from the São José de Itaboraí Basin. It belongs to a bird of the size of Itaboravis elaphrocnemoides and its distinctive morphology does not correspond to any living taxon.
Tambussi and Degrange ( 2013: 31) mentioned a tibiotarsus possibly belonging to Itaboravis (probably referring to this specimen), although Mayr et al. only referred to similar dimensions.
210. †Neornithes indet. 6
Mayr et al. ( 2011a ) reported the distal end of a left tibiotarsus (MN 4117-V) from the São José de Itaboraí Basin. It shares similarities with tinamids, but its fragmentary state does not allow a reliable determination.
211. †Neornithes indet. 7
Mayr et al. ( 2011a ) reported the distal end of a left tibiotarsus (MN 4118-V) from São José de Itaboraí Basin. It is poorly preserved, with no distinctive characters.
212. †Neornithes indet. 8
Early Eocene – RJ
Aves indet. (?D or E) – Mayr et al. 2011a: 683, 2E.
Mayr et al. ( 2011a ) reported the fragmentary distal end of a left tarsometatarsus (MN 4120-V) from the São José de Itaboraí Basin. It consists of the trochlea metatarsi III only, with a size that approximately matches that of the two tibiotarsi mentioned above.
213. †Neornithes indet. 9
Late Miocene – AM
“fragmento de ulna de ave” – Ramos et al. 2001: 172.
Ramos et al. ( 2001 ) reported an ulna fragment from the Torre da Lua outcrop of Solimões Formation at the margins of the Tarauacá River near Eirunepé, Amazonas.
214. †Neornithes indet. 10
Souza-Filho and Guilherme ( 2015 ) reported avian material from the Morro do Careca site in the BR-364 highway between Feijó and Tarauacá, Acre.
215. Neornithes indet. 11 (spp.)
Souza Cunha ( 1962 ) reported relatively large tarsometatarsi from the Olho d’Água da Escada site in Baraúna (then part of Mossoró), Rio Grande do Norte, collected in February 1961 in association with several mammalian bones. Subsequent analysis of these sediments revealed many avian bones associated with microvertebrate remains. The material is well preserved and shows various degrees of fossilization, from completely mineralized to fresh, recent-looking bones (most of the remains). It is represented mostly by limb bones, synsacra, and vertebrae, and includes various taxa. These comprise the first record of pleistocenic avian remains from Rio Grande do Norte.
216. Neornithes indet. 12 (spp.?)
Pleistocene – ES
Aves – Paula Couto 1978a: 378.
Paula Couto ( 1978a ) reported pleistocenic long bone fragments of small birds from a quarry in Serra da Gironda in Cachoeiro de Itapemirim, Espírito Santo. The material was collected in 1974 by Rubens da Silva Santos and is deposited in the MN. Besides these avian remains, bones of mammals (including megafauna), reptiles, and amphibians were discovered in the same deposit.
217. Neornithes indet. 13
Pleistocene – RO
“um fêmur esquerdo de ave de espécie ainda não identificada” – Bissaro Júnior et al. 2009: 157.
Bissaro Júnior et al. ( 2009 ) reported an indeterminate left femur from the left bank of the Madeira River around the construction area of the Santo Antônio Hydroelectric Dam in Porto Velho, Rondônia. Although lacking stratigraphic context, the bone is permineralized and shows no evidence of intense transport, being found near outcrops related to the Rio Madeira Formation (27,310±200 to >46,310 years BP).
218. Neornithes indet. 14
Hsiou ( 2009a ) reported a tibiotarsus (MCN-PV 8806) from the Touro Passo Formation ( Fig. 1 .15) in Uruguaiana, Rio Grande do Sul. Its high fragmentation and fragility prevented an attribution even to the family level.
219. Neornithes indet. 15 (spp.)
Scherer et al. ( 2012 ) reported several avian remains from the Pleistocene–Holocene of Toca dos Ossos in Ourolândia, Bahia. The material comprises 43 specimens: two fragmentary mandibles, seven humeri, a radius, three femora, seven tibiotarsi, six tarsometatarsi, and 17 indeterminate fragments.
220. Neornithes indet. 16 (spp.?)
From Picos II, a lagoon deposit in the Fazenda Picos in Piranhas, Alagoas, Oliveira et al. ( 2013 ) reported fragments of quaternary-age avian bones associated with megafaunal remains. They figure among material collected in January 2011 and January 2012, though Silva et al. ( 2013 ) mentioned a bird bone from that site collected in 2010. Nevertheless, this is the first record of bird bones for this kind of deposit in Alagoas. The fragments are small and could not be determined at a more specific level. The largest fragment (SGP-MHN-UFAL 0931-V) is part of a diaphysis of a long bone 70 mm long with a diameter of 10 mm and preserved trabeculae, similar to a femur, a tibiotarsus, or a tarsometatarsus of an aquatic bird or a bird of prey.
221. Neornithes indet. 17
Castro et al. ( 2014 ) provisionally attributed to a charadriiform or a passeriform a tibiotarsus from the Pleistocene of Gruta do Ioiô in Palmeiras, Bahia.
222. Neornithes indet. 18 (spp.)
Faure and Guérin reported several small bones (PISF-18-637-10, PISF-18-972, PISF-18-5335, and PISF-18-5692) and three fairly large long bones (PISF-18-672 [incomplete], PISF-18-3156, and PISF-18-7899) of late pleistocenic age from the Lagoa Uri de Cima site in Salgueiro, Pernambuco. The material was found during 2010–2013 digging campaigns as part of the Projeto de Integração do Rio São Francisco com as Bacias Hidrográficas do Nordeste Setentrional (PISF) and is deposited in the Laboratório de Paleontologia of FUMDHAM. Mayer ( 2013 ) reported cathartid remains from the same site that possibly belong to this material.
223. Neornithes indet. 19
Patusco et al. ( 2016a , 2016b ) reported the distal end of a left tarsometatarsus (MN 2874-V) from tank 3 of the João Cativo locality in Itapipoca, Ceará.
224. Neornithes indet. 20
Quaternary – CE
“aves” [in part] – Patusco et al. 2016b: 274.
Patusco et al. ( 2016b ) reported a large phalanx (MN 2993-V) from tank 3 of the João Cativo locality in Itapipoca, Ceará.
Palaeognathae Pycraft
† Diogenornis Alvarenga
225. † Diogenornis fragilis Alvarenga
Early Eocene – RJ
Fig. 31 A
Type locality: Itaboraí limestone Basin, about seven hundred meters east of the São José village, district of Cabuçu, Municipality of Itaboraí, State of Rio de Janeiro, Brazil.
Etymology:Diogenornis , with the Greek ornis , “bird of Diógenes”, honoring Brazilian paleontologist Diógenes de Almeida Campos (1943–) (DNPM) for the support given to Alvarenga, and fragilis , Latin for “fragile”, referring to the idea that the bird was vulnerable to predators because of its terrestrial habit.
Aves [“Ainda por determinar”; in part?] – Paula Couto 1958a: 11.
Aves [“Diversas, não identificadas”; in part?] – Paula Couto 1970: 909.
Aves [“fragmentos de ossos longos”; in part?] – Palma 1973: 40.
Aves [“fragmentos de ossos longos”; in part?] – Palma and Brito 1974: 400.
“restos de diversas aves, metatarsais, vértebras, pré-maxilas, etc.” [in part?] – Price (in lit. 1977 in Sick 1984a ): 65.
Aves [“representadas por ossos longos”; in part?] – Francisco and Souza Cunha 1978: 390, 402, 408.
Diogenornis fragilis , sp. n. [new genus and species] – Alvarenga 1983a: 1–8, figs 1–17.
Diogenornis fragilis – Mones 1986: 75.
Diogenornis fragilis – Cuello 1988: 4.
Diagenornis [lapsus] – Peters 1988: 226.
“fragmentos de ossos longos de aves” [in part?] – Brito 1989: 60.
Diogenornis fragilis – Alvarenga 1993a: 62, 33.
Diogenornis fragilis – Alvarenga 1993b: 22, 14.
Diogenornis fragilis – Unwin 1993: 720.
Diogenornis fragilis – Alvarenga 1997: 123, 33.
Diogenornis fragilis – Kellner 1998: 654, 658–659.
Diogenornis – Kellner and Campos 1999: 247, 9.
Diogenornis fragilis – Alvarenga and Höfling 2000: 587.
Diogenornis fragilis – Alvarenga and Höfling 2004: 825.
Diogenornis fragilis – Bergqvist et al. 2006: 55, 62.
Diogenornis fragilis – Bergqvist et al. 2009: 422, 9G.
Diogenornis fragilis – Mayr 2009: 32, 5.2A–G.
Diogenornis fragilis – Alvarenga and Höfling 2011: 123.
Diogenornis fragilis – Mayr 2022: 38–39, 3.5A.
Alvarenga ( 1983a ) described this new genus and species from sparse and fragmented bones from the São José de Itaboraí Basin, housed in the DNPM (now MCT) and MN collections. The DNPM material, most of which was collected in 1949 by Júlio da Silva Carvalho, was not examined in detail until Déa Regina Bouret Campos (then responsible for the institution’s paleontological collection), aware of Alvarenga’s interest in fossil birds, invited him to study it. Later, paleontologist Fausto Luiz de Souza Cunha from MN sent Alvarenga fragments of bird bones with the same origin collected in 1948 by naturalist Ney Vidal, from the former Divisão de Geologia of MN. After preparing and restoring several bones, Alvarenga concluded that most belong to a single species and represent 4 or 5 individuals.
The holotype comprises the complete left and right tibiotarsi and a right tarsometatarsus lacking the distal end of a young specimen (MCT.R.1421). The paratypes consist of a right tarsometatarsus of an adult individual, lacking the diaphysis’ distal half but retaining the distal extremity (MCT.R.1422); a left tarsometatarsus of a young adult lacking the diaphysis’ distal shaft, also preserving the distal extremity (MCT.R.1423); the proximal end of a right tarsometatarsus (MN 4033-V); the middle segment of the diaphysis of a right tarsometatarsus (MN 4034-V); a segment corresponding to the distal metaphysis of a left tarsometatarsus, lacking the lateral part (MN 4035-V); the middle trochlea of a left tarsometatarsus (MN 4036-V); the proximal end of a left humerus (MCT.R.1424), which appears to be larger than that of Rhea (although Diogenornis is overall much smaller); the proximal phalanx of the major digit of the left wing (MCT.R.1430), narrower and longer than in Rhea ; anterior part of the premaxilla (MCT.R.1428), showing a robust, narrow beak very different from that of the living rheas, resembling that of a chicken, or even a cassowary, as noted by Olson ( 1985b: 106); posterior part of the vertebral bodies of five cervical vertebrae (MCT.R.1425), probably C-5 to C-9; nine cervical vertebrae, probably associated, including atlas, axis, and others identified as C-3, C-4, C-5, C-7, C-9, C-10, and C-11 (MCT.R.1426); six thoracic vertebrae (MCT.R.1427), last one probably pre-synsacral; axis (MCT.R.1429); thoracic vertebra (MN 4037-V), probably the first or second, and another thoracic vertebra (MN 4038-V), probably the pre-synsacral penultimate or antepenultimate.
The bird was probably flightless, but its wings were proportionally much more developed than those of Rhea , which raises the possibility that its close ancestors were flighted (Alvarenga 1983a ). Its dimensions were estimated between 80 and 90 cm in height (Alvarenga 1983a ) and a weight of 4.138 kg (Taranto and Bergqvist 2010 ) or approximately 11 kg (Crouch and Clarke 2019 ). Furthermore, Taranto et al. ( 2011 ), while studying the bird’s hind limbs, estimated the length of the absent femur and concluded that the legs’ morphology is comparable to that of Rhea americana , but with the dimensions of a young specimen, which suggested a possible congruence in the biomechanical locomotion between the genera. This topic was addressed in greater detail by Taranto ( 2012 ), who pointed out that Diogenornis , being a cursorial, digitigrade bird with three large and robust toes, would be well adapted to running and could achieve a speed close to the 50 km/h that Rhea can reach [23] .
The discovery of several individuals of different sizes and ages together may indicate sexual dimorphism and gregarious habits (Alvarenga 1983a ).
Alvarenga ( 1983a ) initially classified the species within the Opisthodactylidae as a probable representative of the ancestors of living rheas and highlighted the beak shape as an essential differentiating character between this family and Rheidae. Houde ( 1988 ) followed this classification and noted close similarities with the European ratite genus Palaelotis and cassowaries. Peters ( 1988 ) and Peters and Storch ( 1993 ) also noted similarities with Palaelotis . Tambussi ( 1995 ) associated it with the Rheiformes and considered the species, along with rheid [24] proximal phalanges from Las Flores (Grupo Río Chico in Chubut, Argentina; early Eocene [Woodburne et al. 2014b ]), as the oldest records for the order. She also concluded that the Rheidae would have been represented since its earliest records by two morphotypes, one including the slender forms of the Eocene of Brazil and the other with a structural pattern present in living species, such as the bird of Las Flores. On its classification as an opisthodactylid, Mayr ( 2009 ) noted that the similarities are probably plesiomorphic and preferred to treat it as a stem rheid. He also pointed out similarities with palaeotidids and remiornithids (only the latter mentioned in Mayr 2022 ). Later, Mayr ( 2016 ) added that there is no solid basis for an affinity between Diogenornis and Rheiformes, but this is very likely due to biogeographic reasons and the general similarity to the living rheas, as also pointed out by Agnolín ( 2016b ).
Alvarenga ( 2010 ) considered it much closer to the Australian casuariids than the South American rheids after restudying the material and comparing it with all ratite families. This reinforces the theory of a Gondwanan origin for these birds, the close relatedness between rheids and casuariids, and the importance of the transantarctic biotic exchange between South America and Australia in the earliest Cenozoic. This possible relation was earlier briefly mentioned by Alvarenga and Höfling ( 2000 , 2004 ). In contrast, a phylogenetic study conducted by Yonezawa et al. ( 2017 ) found it to be closer to the rheas. Picasso et al. ( 2022 ) stated the immature condition and fragmentary nature of the available specimens constitute a strong motivation to further comparisons including casuariids and rheids of different ontogenetic stages to reevaluate its taxonomic status.
Agnolín and Cenizo ( 2014 ) tentatively kept Diogenornis as a casuariid and assigned to the genus the distal end of a tibiotarsus from the Río Chico locality (Sarmiento Formation, middle Eocene–lower Miocene levels; Picasso et al. 2022 ) in Chubut, Argentina. They also mentioned the distal end of another tibiotarsus from the Bryn Gwyn locality (Sarmiento Formation, early Oligocene levels; Picasso et al. 2022 ) in Chubut, which is similar to Diogenornis and the casuariids. Later, Agnolín ( 2016a ) referred to the Río Chico tibiotarsus as cf. Diogenornis within the Ratitae and pointed out similarities with the casuariids, and the tibiotarsus from Bryn Gwyn was left as indeterminate Aves after a detailed review due to its fragmentary nature (F.L. Agnolín, personal communication). In the face of the new hypothesis of a relationship between the oldest South American ratites and the casuariids, Agnolín ( 2016a ) analyzed unpublished specimens deposited in several Argentinean institutions and reevaluated published specimens from different sites in the country. His results suggested that, during the Paleogene and the beginning of the Neogene, the South American ratites were quite diverse, encompassing not only rheids but also several taxa of uncertain affinity that were possibly extinct with the replacement of their warm forested habitats by arid areas at the end of the Miocene. This unexpected ratite diversity in South America and the equivalent scenario in Europe and Africa, along with recent molecular analyses, made him question the model of the modern ratites’ origin as exclusive products of the breakup of Gondwana. Mayr ( 2022 ) challenged this hypothesis of a higher diversity of non-rheiform paleognathous birds and noted that the similarities between rheas and other large flightless birds are due to convergence (based on the results of current molecular analyses) and archaic stem-group rheiforms are expected to be very distinct from more advanced taxa.
Despite close affinities between South American and Australian taxa being possible from a biogeographic perspective, Mayr ( 2022 ) commented that the Casuariidae and its sister taxon Dromaiidae are very similar regarding the morphology of their postcranial skeleton and Diogenornis differs from both in plesiomorphic skeletal traits, with comparisons with other Paleogene paleognathous birds needed for informed assessment of its affinities.
Metello et al. ( 2012a ) associated with this species an ungual phalanx (UFRJ-DG 305 M) with the same origin as the other known elements, which was erroneously deposited in the Mammal Collection of the Departamento de Geologia of UFRJ.
Metello ( 2013 ) mentioned two ungual phalanges attributed to this species and noted they consist of digits III and IV, probably of the left foot. One is the above record, and the other is specimen MCT 1839-R (Metello and Bergqvist 2014 ). The phalanges are similar to those of Struthio , probably due to the basal position of Diogenornis , and their analysis reinforces the cursorial habits attributed to the taxon (Metello et al. 2012a , Metello 2013 ). Metello and Bergqvist ( 2014 ) evaluated the curvature of the phalanges and interpreted it as a well-established terrestrial bird (even though it still retained some reduced flight capacity), that possibly behaved like a tinamid, foraging and nursing in the ground and flying short distances to escape predators.
Rheiformes Forbes
Rheidae Bonaparte
Rhea Brisson
226. cf. Rhea sp.
Quaternary – CE
Aves incertae sedis – Araújo Júnior 2012: 70.
Struthioniformes – Metello and Araújo Júnior 2012: 18.
Struthioniformes – Metello and Araújo Júnior 2013: 70.
Aves incertae sedis – Araújo Júnior et al. 2013: 57.
Aves incertae sedis – Araújo Júnior 2015: 191.
Rhea ? [?; in part?] – Araújo Júnior 2016: 150.
“a large bird” – Asakura et al. 2016: 164.
Rhea ? [?; in part?] – Waldherr et al. 2017: 471.
Rhea ? [?; in part?] – Waldherr et al. 2019: 115.
Araújo Júnior ( 2012: 70, 110) and Araújo Júnior et al. ( 2013 ) reported a long bone fragment of a large indeterminate adult bird from the Jirau site in Itapipoca, Ceará. They noted that only Rhea was recorded in the natural tanks, but the state of the material precluded a more detailed description. It was thus determined only as Aves incertae sedis. It is noteworthy that material from Piauí was assigned to Rhea fossilis (Faure et al. 2010b ; but read on). The fossil is deposited in the MUPHI collection.
227. Rhea sp.
Rock paintings of the Nordeste Tradition interpreted as rheas exist in rock shelters of the Parque Nacional Serra da Capivara, such as those pictured grouped in a row from Toca do Boqueirão da Pedra Furada, Toca da Pinga do Boi, Baixão do Perna I, Toca do Boqueirão do Puxa, and Toca do Vento (Guidon 1991 , Pessis 2003 , Faure et al. 2010b ). However, despite the rich vertebrate fauna excavated in the region since the 1980s, surprisingly no remains of these birds were found (Guérin et al. 1993a , 1993b , 1996 ). This changed when Guidon et al. ( 2009b ) first reported the genus Rhea from Toca do Serrote das Moendas, a rock shelter that started being excavated in 2006.
Faure et al. ( 2010a , 2010b ) attributed the material, the distal end of a left tarsometatarsus (labeled as 113–145606) from layer 15 of sector 1 of that site to Rhea fossilis (Moreno and Mercerat 1891 ), a taxon based on fragments of tibiotarsus and two tarsometatarsi from the quaternary Luján Formation of Mar del Plata, Buenos Aires Province (Picasso et al. 2022 ), that was similar but slenderer than Rhea americana . The bone is incompletely preserved and bears some markings of rodent teeth. Remains of pampatheriines, dasypodids, glyptodontids, scelidotheriines, macraucheniids, equids, cervids, tayassuids, camelids, canids, felids, small rodents, various reptiles, and amphibians were also found in the same layer.
Reevaluations of the original material of Rhea fossilis and other Argentinean quaternary paleospecies ( Rhea anchorenensis , Rhea pampeana , and Rhea subpampeana ) considered them all to be invalid, being instead representatives of the living Rhea americana (Tonni 1980 , Picasso et al. 2022 ). Since the Brazilian material was not included in that analysis, it will be treated here provisionally as Rhea sp.
228. cf. Rhea americana (Linnaeus)
Holocene – SE
“possível Ema ( Rhea americana )” – Oliveira et al. 2016a: 4.
Oliveira et al. ( 2016a ) reported 29 necklace beads possibly made of rhea long bones in a funeral adornment found in human burial 112 of Justino site in the Xingó sub-region of Canindé de São Francisco, Sergipe. The material is deposited in the Laboratório de Bioarqueologia (LABIARQ) collection of UFS.
229. Rhea americana (Linnaeus)
Quaternary – CE, GO, MG, MS, PE, RS, SP
Figs 6C, 18A–C
“deux espèces d’Autruches ( Rhea ) dont l’une bien plus grande que l’espèce actuelle” [in part] – Lund 1840: 319.
“uddöd Art af Familien Alectorides Ill. af Störrelse som den amerikanske Struds” [in part?] – Lund 1841d: 18.
“deux espèces d’autruche, dont l’une bien plus grande que l’espèce actuelle” [in part] – Claussen 1841: 20.
“to Arter Struds, hvoraf den ene betydelig större end den her nulevende Art” [in part] – Lund 1842b: 130.
“un genre voisin des Dicholophus ou Cariamas” [?] – Gervais 1844a: 294.
“Autruche du genre Rhea , plus grande que les deux espèces actuelles” – Gervais 1844b: 33.
“l’une des plus remarquables par sa grande taille est de la famille des Alectoridés ou Hoccos” – Gervais 1844b: 34.
Dicholophus [in part] – Giebel 1846: 311.
Rhea [?; in part] – Giebel 1846: 312.
Rhea [in part] – Giebel 1847: 24.
Dicholophus [in part] – Giebel 1847: 27.
“deux espèces à trois doigts (sous-genre Rhea ) dont une est bien plus grande que celle qui vit aujourd’hui dans l’Amérique méridionale” [in part] – Pictet 1853: 416.
“une espèce plus grande que les espèces vivantes. Mais celle-ci n’appartient pas à un nouveau genre. Elle était du type du Nandou, ou autruche du Brésil, et appartenait comme elle au genre Rhea” – Liais 1872: 302.
“le genre Cariama ( Dicholophus d’Illiger, Microdactylus de Geoffroy, Seriema du Brésil)” [?] – Liais 1872: 303.
“two species of American ostrich ( Rhea ), one larger than either of the living species” [in part] – Wallace 1876: 164.
“gigantisk Styltegænger” – Lund (in Reinhardt 1881 ): 147.
Rhea americana [in part] – Reinhardt 1881: 141–153, figs 1–3.
Rhea americana (et aff.) [in part] – Winge 1887: 18.
Rhea americana [in part] – Goeldi 1894: 614.
Rhea americana [in part] – Lambrecht 1921: 3.
Rhea americana [in part] – Lambrecht 1933: 729.
Rhea americana [in part] – Brodkorb 1963: 201.
Rhea americana [in part] – Mones 1986: 75.
Rhea americana [in part] – Cuello 1988: 26.
Rhea americana [in part] – Nascimento and Silveira 2020: 489.
Lund ( 1840 , 1842b ) identified remains of Rhea americana among his finds in the Lagoa Santa region, interpreting them as two species, one larger than the living one. Later, in the summary of his unpublished treatise (Lund 1841d ), he mentioned only one species for the genus and one form he believed to be an extinct, Rhea -sized alectorid. Reinhardt ( 1881 ) noted that Lund, in his unpublished treatise, apparently did not believe he had the remains of more than one species of Rhea , and from that, it can be concluded, although not explained, that the alectorid is the same bird as one of these species of rhea.
Several subsequent authors repeated this treatment as two species of rheas represented in the material (e.g., Claussen 1841 , Wallace 1876 ). Gervais ( 1844b ) mentioned the two rhea species and the large alectorid. Liais ( 1872: 302) mentioned the two rhea species and “le genre Cariama ( Dicholophus d’Illiger, Microdactylus de Geoffroy, Seriema du Brésil)”, noting the latter had a species in quaternary times. He was probably referring to Gervais’ account of “un genre voisin des Dicholophus ou Cariamas” supposedly found by Claussen, whose alleged finds Liais attributed to Lund. Winge ( 1887 ) stated he did not know what this designation referred to.
The bones initially determined as the alectorid are the proximal end of a right tarsometatarsus and a pedal phalanx lacking the proximal joint surface, both impregnated with incrustations (Reinhardt 1881 ). Reinhardt reported that, according to Lund’s own words in his unpublished treatise, both fragments were found in the same place in the same cave, Lapa da Anna Felicia (Winge 1887 ), and had all the markings that indicated that they belonged to the same individual. Colored illustrations of both (done by Peter Andreas Brandt) were included in this treatise (with monochrome versions reproduced in Reinhardt 1881 , 1882 ).
The alectorid was based on these two fragments. However, after sending his treatise to Denmark, Lund must have realized that he had a third bone of that bird: the middle part of a right tibiotarsus covered in incrustations, which was marked in his catalog (erroneously as a femur) as No. 8 and coming from the same cave as the other two fragments, under the name of “gigantisk Styltegænger” (“giant wader”), the same under which the tarsometatarsus, of No. 9, was cataloged (Reinhardt 1881 ). Reinhardt noted that both correspond in form, appearance, and condition, and it can be concluded they are the remains of the same individual and that Lund does not appear to have found more than these three fragments of this bird since this name does not appear again in his catalog, nor any mention of Alectorides.
Material from tank 2 of the João Cativo site. Rhea americana: distal end of the right tarsometatarsus (MN 3275-V), anterior (A), caudal (B), and medial (C) views. Accipitridae indet. 9: ungual phalanx (MN 3263-V), medial (G), lateral (H), and proximal (I) views (H.I. Araújo Júnior; C.L. Ximenes). Scale bar: 10 mm.
According to Reinhardt, Lund thought the giant alectorid was generically different from Cariama —the only component of this unnatural group with which Lund was more familiar and could compare bones—and was the missing link between it and Palamedea . This opinion was based on the differences observed between the tarsometatarsi [25] and, perhaps mainly, on the pedal phalanx, which would indicate longer toes than the seriemas. However, unfortunately, this bone seems to have been lost or destroyed still in Brazil since there is no mention of it in Lund’s catalog, and not even Reinhardt was able to locate it in the collection.
The thorough examination of the fragments and the illustration of the pedal phalanx led Reinhardt to conclude that these bones belonged to an adult specimen of Rhea americana . On the possibility of dealing with another species of the genus, he noted that, although he has not recognized anything that goes against his conclusion, the discovery of new bones can bring out differences that could not be inferred from such limited material.
While agreeing with Reinhardt’s conclusion about the identity of the tibiotarsus and tarsometatarsus fragments, Winge ( 1887 ) considered that the pedal phalanx is most likely the first phalanx of a llama, and the lack of its proximal part contributed to the confusion. He also noted Lund might have realized this, and perhaps this is the reason for the absence of this bone in his catalog.
“deux espèces d’Autruches ( Rhea ) dont l’une bien plus grande que l’espèce actuelle” [in part] – Lund 1840: 319.
“ Rhea ” – Lund 1841d: 18.
“deux espèces d’autruche, dont l’une bien plus grande que l’espèce actuelle” [in part] – Claussen 1841: 20.
“to Arter Struds, hvoraf den ene betydelig större end den her nulevende Art” [in part] – Lund 1842b: 130.
Rhea – Gervais 1844b: 34.
Rhea [?; in part] – Giebel 1846: 312.
Rhea [in part] – Giebel 1847: 24.
“deux espèces à trois doigts (sous-genre Rhea ) dont une est bien plus grande que celle qui vit aujourd’hui dans l’Amérique méridionale” [in part] – Pictet 1853: 416.
“Outre cette grande espèce, le Dr Lund en a encore trouvé une seconde du même genre, plus voisine de l’espèce actuelle par sa taille” – Liais 1872: 302.
“two species of American ostrich ( Rhea ), one larger than either of the living species” [in part] – Wallace 1876: 164.
Rhea – Reinhardt 1881: 142.
Rhea [in part] – Lund (in Winge 1887 ): 18.
Rhea aff. americanæ [in part] – Lund (in Winge 1887 ): 18.
Rhea americana (et aff.) [in part] – Winge 1887: 18–19.
Rhea americana [in part] – Goeldi 1894: 614.
Rhea americana [in part] – Lambrecht 1921: 3.
Rhea americana [in part] – Lambrecht 1933: 729.
Rhea americana [in part] – Brodkorb 1963: 201.
Rhea americana [in part] – Mones 1986: 75.
Rhea americana [in part] – Cuello 1988: 26.
Rhea americana [in part] – Nascimento and Silveira 2020: 489.
From the material of Rhea americana that was not primarily determined as the alectorid, Winge ( 1887 ) reported a second phalanx of digit III (“ Rhea ” in Lund’s catalog) from Lapa da Anta I, slightly smaller than in R. americana , and from Lapa da Escrivânia I several bones: a sixth cervical vertebra lacking the anterior part, the distal end of a tibiotarsus (“ Rhea aff. americanae ” in Lund’s catalog) with slight variation in morphology, the fragment of a distal end of a tarsometatarsus (“ Rhea aff. americanae ” in Lund’s catalog), and a first phalanx of digit II, all smaller than in R. americana . Regarding the possibility that these smaller bones from Lapa da Anta and Lapa da Escrivânia I represented a different species, Winge noted that it is very unlikely, with no reason to think about any other species besides the living one.
“Neornithes: indeterminadas” [in part] – Souza Cunha 1961: 5.
“Neornithes, Ordens e gêneros indeterminados” [in part] – Paula Couto 1961: 8.
“aves (indeterminadas)” [in part] – Paula Couto 1962: XIX.
“Aves Neornithes” [in part] – Paula Couto 1978b: 429.
Rhea – Paula Couto 1980: 145.
“Aves neornites” [in part] – Silva Santos 1982: 118.
Rhea – Ximenes 2009: 472.
Rhea – Souto and Carvalho 2010: 116.
Rhea sp. – Araújo Júnior 2012: 70.
“gavião fóssil” [in part] – Metello and Araújo Júnior 2012: 18.
Rhea sp. – Araújo Júnior et al. 2013: 59.
“gavião fóssil” [in part] – Metello and Araújo Júnior 2013: 69–70.
Acciptriformes indet. [sic] [?; in part] – Araújo Júnior 2015: 193.
Rhea ? – Araújo Júnior 2016: 150.
Rhea sp. – Asakura et al. 2016: 164.
“bird remains” [in part] – Patusco et al. 2016a: 190.
“aves” [in part] – Patusco et al. 2016b: 274.
Rhea ? – Waldherr et al. 2017: 471.
Rhea ? – Waldherr et al. 2019: 115.
Rhea americana – Costa et al. 2024: 3, 2K–L.
Paula Couto ( 1980 ) reported the genus Rhea among the quaternary fauna discovered in tank 2 of the João Cativo locality in Itapipoca, Ceará. From that material, Costa et al. ( 2024 ) assigned to Rhea americana the distal fragment of a right tarsometatarsus (MN 3275-V), tentatively attributed to a juvenile or sub-adult individual. It was previously mentioned among remains associated with an accipitrid (see Accipitridae indet. 9).
Patusco et al. ( 2016a , 2016b ) reported the proximal end of a right tarsometatarsus (MN 3269-V) that possibly represents the same bone as specimen MN 3275-V.
“emas” – Collet and Prous 1977: 33.
Collet and Prous ( 1977 ) mentioned rhea bones among the remains from fluvial sambaquis in Itaoca, São Paulo. The presence of these bones in these archeological sites suggests human contact with the neighboring plateau.
Rhea americana [in part] – Schorr 1976: 98.
Rhea americana [in part?] – Schmitz 1980: 200.
Rhea americana [in part] – Schmitz et al. 1981: 93.
Rhea sp. [in part] – Jacobus and Schmitz 1983a: 58, 9.
Rhea sp. [in part] – Jacobus and Schmitz 1983b: 269, 8.
Rhea – Schmitz and Jacobus 1983 –1984: 34.
Rhea – Jacobus 1985: 64.
“ema” [in part] – Jacobus (in Schmitz and Machado 1987 ): 167.
Rhea americana – Schmitz et al. 1989: 154–155.
Rhea americana [in part] – Barbosa et al. 1990: 89.
“emas” [in part?] – Schmitz 1990: 110, 2.
“ema” – Schmitz 1999: 93.
Rhea americana – Dias 2004b: 255.
Rhea americana – Rosa 2004: 230, 233, 236.
Rhea americana – Paulo 2009: 141.
Rhea americana – Da-Gloria and Larsen 2017: 105.
Remains from the GO-JA-01 rock shelter in Serranópolis, Goiás, were mentioned in the literature on a handful of occasions. Schorr ( 1976 ) reported this species among the archeofaunal remains from GO-JA-01, GO-JA-14, and GO-JA-20. Jacobus and Schmitz ( 1983a , 1983b ) reported the genus Rhea from GO-JA-01 and noted the material they analyzed comes from different stratigraphical cuts (III) than that reported by Schorr (I–II). They depicted the distal part of the right tibiotarsus, the proximal part of the right tarsometatarsus, two distal ends of left tarsometatarsi, and a fragment of vertebra, besides several other indeterminate avian remains. Schmitz ( 1990 ) depicted these elements, except for the vertebra, and noted they were modified by humans. Jacobus (in Schmitz and Machado 1987 ) noted the four identified fragments come from the site’s square 12H. Schmitz et al. ( 1989 ) reported that the material (including that analyzed by Jacobus and Schmitz) from GO-JA-01 included the maxilla, mandible, axis, vertebra, sternum, coracoid, humerus, radius, ulna, carpometacarpus, synsacrum, femur, tibiotarsus, and tarsometatarsus, besides more numerous eggshells. Rosa ( 2004 ) reanalyzed this material and reported six bones of at least one individual from the site’s square 12H, two bones of at least one individual from 14H, and a bone from 16H, attributing them to the Paranaíba phase (early Holocene). The material is deposited in the IAP/Unisinos collection.
Goldmeier and Schmitz ( 1983 ) mentioned “ostriches” (= Rhea americana ) among the kitchen remains from cerritos of the Chuí phase of the Umbu tradition in Santa Vitória do Palmar, Rio Grande do Sul, dating back to about 2,400 or 2,500 years BP.
Mentz Ribeiro et al. ( 1989 ) reported this species from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Rosa ( 2009 ) reported the material to comprise eight bones (two from square C6 and six from D6) and attributed it to this species with uncertainty. The material is deposited in the CEPA collection.
“ema” – Lima 1991: 57.
Lima ( 1991 ) reported necklace beads made of rhea and other bird bones discovered in human graves of about 2,000 years BP at Furna do Estrago in Brejo da Madre de Deus, Pernambuco.
Rosa ( 2001 , updated in 2006b ) reported four bones (one from level 2, two from level 3, and one from level 4) from the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. Eggshells were also found. The material is deposited in the IAP/Unisinos collection.
Rhea americana – Santos et al. 2007b: 2.
Santos et al. ( 2007b ) reported this species among the archeofaunal remains from sites in the western border of Pantanal in Mato Grosso do Sul.
Rhea americana – Jacobus and Rosa 2013: 247.
Jacobus and Rosa ( 2013 ) reported remains from the Schneider (RS-C-14) rock shelter in São Sebastião do Caí, Rio Grande do Sul.
Rhea americana – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
Rhea americana – Peixoto and Silva 2017: 16.
Peixoto and Silva ( 2017 ) reported tarsometatarsus epiphyses and phalanges from the late Holocene of the Limoeiro mound in the Corumbá region, Mato Grosso do Sul.
Tinamiformes Huxley
Tinamidae Gray
230. Tinamidae indet. 1
Carvalho ( 1984 ) reported tinamid remains (including a worked femur) among the material discovered in 1978 at the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
231. Tinamidae indet. 2
Late Holocene – SP
“Tinamídeos” – Figuti 1993: 74.
Figuti ( 1993 ) reported tinamid remains from the COSIPA sambaqui 2 in Ilha do Casqueirinho, Cubatão, São Paulo.
232. Tinamidae indet. 3 (spp.)
Holocene – GO
Tinamidae indet. [in part] – Rosa 2004: 230.
Tinamidae indet. [in part] – Rosa 2004: 233.
Tinamidae indet. sp. 1 [in part] – Rosa 2004: 237.
Tinamidae indet. sp. 2 [in part] – Rosa 2004: 237.
Tinamidae indet. [in part] – Rosa 2004: 239.
Tinamidae indet. [in part] – Rosa 2004: 247.
Tinamidae indet. [in part] – Rosa 2004: 249.
Tinamidae indet [?; in part?] – Paulo 2009: 141.
Rosa ( 2004 ) reported several indeterminate tinamid remains from the GO-JA-01 rock shelter in Serranópolis, Goiás. From the Paranaíba phase (early Holocene), a bone from square 12H, 32 bones of at least 11 individuals from 14H, two bones of at least one individual from 16H, another bone from 16H but of a different species, three bones of at least one individual from 18H, six bones of at least two individuals from 18I, and a bone from the cut 1/2. From the Serranópolis phase (early–late Holocene), a bone from 18I. The material is deposited in the IAP/Unisinos collection.
233. Tinamidae indet. 4
Rosa ( 2009 ) reported 54 bones (34 from square A6, two from B5, one from C7, 15 from D6, and two from D7) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Remains from square A6 were detected in the site’s three occupation periods, with some showing cutting marks and burn traces. The material is deposited in the CEPA collection.
234. Tinamidae indet. 5
Early Holocene – SP
“tinamídeos” – Alves 2008: 170.
Alves ( 2008 ) reported tinamid remains from areas II (at least two individuals) and III (at least one individual) of the Capelinha I site in Cajati, São Paulo, aged 9,250±50 years BP.
235. Tinamidae indet. 6
Holocene – RS
Tinamidae – Jacobus and Rosa 2013: 244.
Jacobus and Rosa ( 2013 ) reported tinamid remains from the Sangão (RS-S-327) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
236. Tinamidae indet. 7
Holocene – RS
Tinamidae – Jacobus and Rosa 2013: 245.
Jacobus and Rosa ( 2013 ) reported tinamid remains from the Deobaldino (RS-S-395) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
237. Tinamidae indet. 8
Holocene – RS
Tinamidae – Jacobus and Rosa 2013: 247.
Jacobus and Rosa ( 2013 ) reported tinamid remains from the Schneider (RS-C-14) rock shelter in São Sebastião do Caí.
238. Tinamidae indet. 9
Holocene – RS
Tinamidae – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported tinamid remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
239. Tinamidae indet. 10
Quaternary – RN
Tinamidae indet. – Costa et al. 2023b: 35.
Costa et al. ( 2023b ) reported a fragment of tibiotarsus among material found in two sites (Ravina do Leon and Ravina das Araras) of the Lajedo de Soledade karstic area in Apodi, Rio Grande do Norte.
Tinamus Hermann
240. Tinamus sp.
Gonzalez et al. ( 2007 ) reported this genus among the material from the Guarani site Piracanjuba in Piraju, São Paulo. The material (with at least part of it collected during May 2004) consists of two distal tarsometatarsus epiphyses from the site’s layers I and II.
241. Tinamus cf. solitarius (Vieillot)
Chahud ( 2021 ) reported the distal end of a right tarsometatarsus (ST-TR-5) of an adult bird from Lapa do Santo in Matozinhos, Minas Gerais. It was previously depicted as an indeterminate bird by Chahud et al. ( 2021 ). The material is deposited in the LEEH-IB-USP collection. Chahud noted the record of this forest species may suggest a biome ecotone or a wetter period in the Lagoa Santa region during the Holocene.
242. Tinamus solitarius (Vieillot)
Winge ( 1887 ) reported the proximal and distal ends of a tibiotarsus from Lapa da Escrivânia V, and a coracoid, an ulna, and the distal end of a tibiotarsus from “various caves” [26] . He noted little difference between this material and a recent skeleton, insufficient to assign it to another species. The distal end of the tibiotarsus differs somewhat from that observed in Crypturellus , Nothura , and Rhynchotus , and the other fragments were determined mainly by their size.
Winge followed Reinhardt’s ( 1870 ) survey of the avifauna from the “Brazilian Campos”, where he reported the occurrence of Tinamus major in the Lagoa Santa region. This record was synonymized with Tinamus solitarius by Hellmayr and Conover ( 1942 ) and Rodrigues ( 2008 ), which is supported by the geographic distribution of these species. T. major occurs mainly in the north and part of the midwest Brazil (Sick 1997 , Cabot et al. 2017a ), and T. solitarius occurs in the southeast (Sick 1997 , Cabot et al. 2017b ), although it is now extinct in the Lagoa Santa region (Rodrigues 2008 ). T. solitarius has also been considered conspecific with T. major (Hellmayr and Conover 1942 , Cabot et al. 2017b ). This record, much like all of Lund’s materials, requires revision, and it is thus better referred to as Tinamus solitarius provisionally.
Klamt ( 2005 ) reported this species from the Röpke (RS-JC-56) “B” site in Ibirama, Rio Grande do Sul, dated 470±50 years BP (Ferrasso and Schmitz 2010 ). He cited Mentz Ribeiro ( 1996 ) as the original source for the record, but we could not access this reference.
Tinamus solitarius – Bueno 2007: 34.
Bueno ( 2007 ) mentioned this species among avian remains from sambaquis in the APA (Área de Proteção Ambiental) de Massambaba, coastal Rio de Janeiro.
Crypturellus Brabourne & Chubb
243. cf. Crypturellus sp.
Schmitz and Ferrasso ( 2011 ) reported a burned humerus comparable to this species from Itapiranga I (SC-U-1), a Guarani site about 1,000 years old in Itapiranga, Santa Catarina. The material was discovered in January 1957 and is deposited in the IAP/Unisinos collection.
244. Crypturellus sp. 1
Rosa ( 2004 ) reported six bones from the Paranaíba phase square 16H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
245. Crypturellus sp. 2
Rosa ( 2004 ) reported three bones of at least one individual from the Paranaíba phase cut 2 of the GO-JA-14 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
246. Crypturellus sp. 3
Early Holocene – RS
Rosa ( 2009 ) reported 207 bones (39 from square A6, 10 from A7, seven from B5, nine from B6, five from B7, 17 from C5, 16 from C6, three from C7, 90 from D6, and 11 from D7) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Remains from square A6 were detected in the site’s three occupation periods, with some showing cutting marks and burn traces. The material is deposited in the CEPA collection.
247. Crypturellus sp. 4
Quaternary – BA
Crypturellus sp. – Silva and Cozzuol 2010: 111.
Silva and Cozzuol ( 2010 ) associated with this genus an incomplete tarsometatarsus lacking the proximal epiphysis among material found in the 1980s at Toca da Boa Vista in Campo Formoso, Bahia. The fossil is deposited in the MHNJB/UFMG collection.
248. Crypturellus obsoletus (Temminck)
Quaternary – MG
Crypturus obsoletus – Winge 1887: 16.
[ Tinamus ] obsoletus – Goeldi 1894: 440.
Crypturus obsoletus – Lambrecht 1933: 730.
Crypturellus obsoletus – Brodkorb 1963: 194.
Crypturellus obsoletus [ Crypturus obsoletus ] – Mones 1986: 74.
Crypturellus obsoletus – Cuello 1988: 27.
Crypturellus obsoletus – Nascimento and Silveira 2020: 489.
Winge ( 1887 ) reported remains from various sites. From Lapa do Baú, a femur fragment. From Lapa do Capão Seco, a coracoid and a carpometacarpus of at least two individuals. From Lapa da Escrivânia V, several individuals, represented by the sternum, coracoid, humerus, and tibiotarsus. From Lapa do Marinho II, several bones of a single individual, represented by the coracoid, humerus, femur, and tarsometatarsus. From Lapa do Taquaral III, a humerus (No. 12209 in Lund’s catalog about this cave). Finally, without recorded origin, but similar in appearance with the last humerus, there is a sternum, an ulna, a femur, and a tibiotarsus. Winge noted the humerus’ proximal end is firmly encrusted in the carpometacarpus’ upper part (both on the left side), with the carpal [27] and the first phalanx of the alular digit in their place. There are also several bones from “various caves” [26] and of unknown origin.
249. Crypturellus noctivagus (Wied)
Quaternary – MG, PI, RS
Crypturus noctivagus – Winge 1887: 16.
[ Tinamus ] noctivagus – Goeldi 1894: 440.
Crypturus noctivagus – Lambrecht 1933: 730.
Crypturellus noctivagus – Brodkorb 1963: 194.
Crypturellus noctivagus [ Crypturus noctivagus ] – Mones 1986: 74.
Crypturellus noctivagus – Cuello 1988: 27.
Crypturellus noctivagus – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported bones of at least two individuals from “various caves” [26] consisting of the coracoid, humerus, ulna, carpometacarpus, and femur. He noted that only the humerus and the femur were compared with recent bones, and the other elements probably belong to the same set of bones. There are also two humeri of unknown origin.
Brodkorb ( 1963 ) erroneously listed it as coming from Lapa da Escrivânia, an error repeated by Cuello ( 1988 ).
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Jacobus ( 2004 ) reported remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
250. Crypturellus parvirostris (Wagler)
Quaternary – MG, PI
Crypturus parvirostris – Winge 1887: 17.
[ Tinamus ] parvirostris – Goeldi 1894: 440.
Crypturus parvirostris – Lambrecht 1933: 730.
Crypturellus parvirostris – Brodkorb 1963: 194.
Crypturellus parvirostris [ Crypturus parvirostris ] – Mones 1986: 74.
Crypturellus parvirostris – Cuello 1988: 27.
Crypturellus parvirostris – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported numerous bones from Lapa da Escrivânia V and several individuals of recent age [28] , including skulls. He noted that, although the bones of this species are supposedly quite distinct from those of Crypturellus tataupa , this is often questionable.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least two adults and one young individual are present in the material.
251. Crypturellus tataupa (Temminck)
Quaternary – MG, RS
Crypturus [?; in part] – Lund 1841d: 18.
Crypturus ? [?; in part] – Giebel 1846: 312.
Crypturus [?; in part] – Giebel 1847: 23.
Tinamus [?; in part] – Pictet 1853: 416.
Crypturus [?; in part] – Pictet 1853: 416.
“le genre Tinamou ( Tinamus de Lath., Crypturus d’Illig., le Nhambú du Brésil)” [?] – Liais 1872: 303.
Crypturus [in part] – Lund (in Winge 1887 ): 16.
Crypturus inter obsoletum et tataupam [in part] – Lund (in Winge 1887 ): 17.
Crypturus tataupa – Winge 1887: 16–17.
[ Tinamus ] tataupa – Goeldi 1894: 440.
Crypturus tataupa – Lambrecht 1933: 730.
Crypturellus tataupa – Brodkorb 1963: 194.
Crypturellus tataupa [ Crypturus tataupa ] – Mones 1986: 74.
Crypturellus tataupa – Cuello 1988: 27.
Crypturellus tataupa – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, more bones from Lapa do Marinho II, two left carpometacarpus fragments from Lapa do Capão Seco (which might be from Crypturellus parvirostris ), and a “dissolved” skeleton of recent age^ [28] ^ found in a cave and determined by Lund. There is a humerus from Lapa do Baú (“1138 Crypturus ” in Lund’s catalog) with an intermediate size between C. obsoletus and C. tataupa , but closer to the latter (of which it can be a very large individual). There is also a skeleton of recent age^ [28] ^ determined by Lund as “ Crypturus inter obsoletum et tataupam ”, about which Winge commented that if it is not this species, there is no corresponding contemporary form in the region. As this tinamid record received a designation in Lund’s catalog, it might be what he referred to as belonging to the genus Crypturus (Lund 1841d ), and what Liais ( 1872: 303) called “le genre Tinamou ( Tinamus de Lath., Crypturus d’Illig., le Nhambú du Brésil)” and noted it had a species in quaternary times.
Jacobus ( 2004 ) reported remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
Rhynchotus Spix
252. Rhynchotus rufescens (Temminck)
Quaternary – MG, RS
Rhynchotus rufescens – Winge 1887: 18.
Rhinchotus rufescens [sic] – Goeldi 1894: 440.
Rhynchotus rufescens – Lambrecht 1933: 730.
Rhynchotus rufescens – Brodkorb 1963: 195.
Rhynchotus rufescens – Mones 1986: 75.
Rhynchotus rufescens – Cuello 1988: 27.
Rhynchotus rufescens – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, a humerus from Lapa da Escrivânia IX, two bones from Lapa da Escrivânia XI, and more bones from Lapa da Lagoa do Sumidouro. The material is represented by the coracoid, scapula, humerus, ulna, radius, carpometacarpus, first phalanx of the major digit of the wing, femur, tibiotarsus, and tarsometatarsus.
“Perdigão ( Rhynchotus sp.)” – Klamt 2005: 47.
Klamt ( 2005 ) reported this species from the Röpke (RS-JC-56) “B” site in Ibirama, Rio Grande do Sul, dated 470±50 years BP (Ferrasso and Schmitz 2010 ). He cited Mentz Ribeiro ( 1996 ) as the original source for the record, but we could not access this reference.
Nothura Wagler
253. Nothura minor (Spix)
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, some of unknown origin (two of them probably from Lapa da Lagoa do Sumidouro), and a scapula of recent age [28] . He noted that, of all the medium-sized species of Nothura and Crypturellus included in his study, there are mostly bones of the shoulder girdle and forelimbs. Given the substantial similarity between the genera, he preferred to leave a large number of hindlimb bones as indeterminate.
254. Nothura maculosa (Temminck)
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, an ulna from Lapa da Escrivânia XI, a humerus from Lapa da Lagoa do Sumidouro, and a coracoid of recent age [28] .
Nothura maculosa – Teixeira and Rosa 2001: C00003.
Teixeira and Rosa ( 2001 ) reported a bone from the Rincão (SC-IÇ-06) site in Içara, Santa Catarina. The material is deposited in the IAP/Unisinos collection.
Rosa ( 2001 , updated in 2006b ) reported seven bones (one from level 1, four from level 2, one from level 4, and one from level 5) from the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Nothura maculosa – Rosa 2006d: 265.
Rosa ( 2006d ) reported fragmentary coracoid and tibiotarsus from the late Holocene of the RS-LC-81 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Rosa ( 2009 ) reported 73 bones (30 from square A6, three from A7, two from B5, two from B6, five from B7, seven from C5, three from C6, three from C7, and 18 from D6) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Remains from square A6 were detected in the site’s three occupation periods, with some showing cutting marks and burn traces. The material is deposited in the CEPA collection.
Nothura maculosa – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported this species from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Nothura maculosa – Bueno 2007: 34.
Bueno ( 2007 ) mentioned this species among avian remains from sambaquis in the APA de Massambaba, coastal Rio de Janeiro.
Nothura maculosa – Silva and Cozzuol 2010: 111.
Silva and Cozzuol ( 2010 ) reported an incomplete tarsometatarsus lacking the proximal epiphysis among material found in the 1980s at Toca da Boa Vista in Campo Formoso, Bahia. The fossil is deposited in the MHNJB/UFMG collection.
Taoniscus Gloger
255. Taoniscus cf. nanus (Temminck)
Quaternary – MG
Tinamus sp. [in part] – Lund (in Winge 1887 ): 17.
Nothura ( Taoniscus ) nana (fere certe) – Winge 1887: 17–18.
[ Nothura ] nana – Goeldi 1894: 440.
Nothura ( Taeniscus ) nana [sic] – Lambrecht 1933: 730.
Taoniscus nanus – Brodkorb 1963: 195.
Taoniscus nanus [ Nothura ( Taoniscus ) nana ?] – Mones 1986: 74–75.
Taoniscus nanus – Cuello 1988: 28.
Taoniscus cf. nanus – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, two bones from Lapa da Escrivânia XI, some isolated bones of unknown origin, and bones of several individuals of recent age [28] , of which a humerus was determined as “ Tinamus sp.” by Lund. The material is represented by the notarium, coracoid, sternum, humerus, ulna, radius, carpometacarpus, synsacrum, femur, tibiotarsus, and tarsometatarsus. Although he had no recent bones for comparison and the species was not recorded alive in the Lagoa Santa region by Lund or Reinhardt, Winge determined the material with almost certainty. He commented that, given the finds of recent remains, it is unlikely the species would not occur regularly in the region since the birds that exist in its geographic range generally coincide with those existing in the Lagoa Santa region, and it may not have been found due to its reclusive habits or by sheer coincidence.
Neognathae Pycraft
Anseriformes Linnaeus
Anhimidae Stejneger
256. Anhimidae indet.
Late Holocene – RS
Anhimidae – Milheira et al. 2019: 45.
Milheira et al. ( 2019 ) reported anhimid remains among the material from the sites PSG-02 and PSG-07 of the Pontal da Barra cerrito complex in Pelotas, Rio Grande do Sul. In a biogeographic context, it likely represents Chauna torquata .
† Chaunoides Alvarenga
257. † Chaunoides antiquus Alvarenga
Late Oligocene/Early Miocene – SP
Fig. 34 A
Type locality: Santa Fé Farm, 2 km north of Tremembé, São Paulo State, Brazil (22°30’S, 45°32’W). Montmorillonitic clay, about 4 m below the most superficial level of shales.
Etymology: Chaunoides , with the Latinized suffix oideus , “similar to Chauna ”, and antiquus , Latin for “ancient”.
Anhimidae – Alvarenga 1993b: 24.
Anhimidae – Alvarenga 1997: 123.
Anhimidae – Mourer-Chauviré 1999: 87.
Chaunoides antiquus , new species [new genus and species] – Alvarenga 1999a: 223–230, figs 3–4, 6–10.
Chaunoides antiquus – Alvarenga and Höfling 2000: 589.
Chaunoides antiquus – Alvarenga and Höfling 2004: 827.
Chaunoides antiquus – Alvarenga and Höfling 2011: 127.
Chaunoides antiquus – Carmo et al. 2024: 5, 3G, J.
Alvarenga ( 1999a ) described this new genus and species from fossils collected between 1978 and 1993 in the montmorillonite clays of the Tremembé Formation in the Fazenda Santa Fé, about 4 m below the shallowest shales. It is the first fossil anhimid to be named.
Remains of at least three individuals are known, which makes it the best-represented (or most abundant) bird in the Tremembé Formation (Alvarenga and Höfling 2011 ). The material consists of an almost complete left coracoid (MN 4619-V, holotype), another incomplete left coracoid (MN 4620-V) associated with an almost complete left femur (MN 4621-V), the distal end of a left ulna (MN 4622-V), the distal end of a right radius (MN 4623-V), a left radius lacking the proximal end (MN 4624-V), a left ulna lacking the proximal end (MN 4632-V) associated with a segment of the distal axis of a left tibiotarsus (MN 4631-V), two segments of the distal end of left tibiotarsi (MN 4625-V and MN 4629-V), the latter associated with the proximal end of a left tarsometatarsus (MN 4630-V), and three unassociated segments of distal axes of right tibiotarsi (MN 4626-V, MN 4627-V, and MN 4628-V). It was smaller and more gracile than the smallest living species in the family, Chauna chavaria , with slender leg bones and less pneumatized skeleton than the living anhimids.
Upon its description, it was the only described fossil species safely associated with the Anhimidae. Recently, Agnolín ( 2022 ) described Chainkanas koshon from a right coracoid with an incomplete distal end unearthed at the Pinturas Formation (early–middle Miocene) of the Santa Cruz province in Argentina. This large species shows closer similarities with Chaunoides than to the extant Anhima and Chauna .
Anatidae Leach
258. cf. Anatidae
Holocene – MS
“uma certa quantidade de ossos de um provável anatídeo” – Schmitz 2002: 207.
Schmitz ( 2002 ) reported a few bones that possible belong to an anatid from some layers of a site in the Jacadigo lagoon in Corumbá, Mato Grosso do Sul. The material was discovered as part of the joint Projeto Corumbá (1990–2001) of IAP/Unisinos and UFMS.
259. †Anatidae indet. 1
Late Oligocene/Early Miocene – SP
Anatidae – Alvarenga (in Castro et al. 1988b ): 2360.
Anatidae – Castro et al. 1988b: 2362.
“fósseis ainda não descritos de Anatidae” – Alvarenga and Höfling 2000: 589.
“fósseis ainda não descritos de Anatidae” – Alvarenga and Höfling 2004: 827.
“fósseis ainda não descritos de Anatidae” – Alvarenga and Höfling 2011: 127.
Anatidae? gen. sp. – Carmo et al. 2024: 5.
Alvarenga (in Castro et al. 1988b) and Alvarenga and Höfling ( 2000 , 2004 , 2011 ) mentioned undescribed anatid remains from the Tremembé Formation. These mentions seem to have changed to podicipedid remains in further studies (Alvarenga 1993b , 1997 ).
260. Anatidae indet. 2 (spp.?)
Figuti ( 1993 ) reported anatid remains from the COSIPA sambaquis 1, 2, and 4 in Ilha do Casqueirinho, Cubatão, São Paulo. Later, he noted (Figuti 1994 –1995) they occurred in large numbers in COSIPA 4, whereas bird remains in the other sites generally occurred in smaller numbers.
261. Anatidae indet. 3
Late Holocene – SC
“pato” – Schmitz and Verardi 1996: 143.
Schmitz and Verardi ( 1996 ) reported anatid remains from the Cabeçudas site in Itajaí, Santa Catarina.
262. Anatidae indet. 4
Holocene – RS
Anatiedae [sic] – Dias and Jacobus 2003: 51.
Dias and Jacobus ( 2003 ) reported anatid remains from the Sangão (RS-S-327) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
263. Anatidae indet. 5
Jacobus ( 2004 ) reported anatid remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
264. Anatidae indet. 6
Late Holocene – RS
Anatidae – Rosa 2006b: 231.
Rosa ( 2006b ) reported two bones (one from level 1 and one from level 4) from the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
265. Anatidae indet. 7
Late Holocene – RS
Anatidae – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported this family from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
266. Anatidae indet. 8
Late Holocene – MS
Anatidae – Schmitz et al. 2009: 347.
Schmitz et al. ( 2009 ) reported two anatid bones from level 3 of the MS-MA-16C site in Corumbá, Mato Grosso do Sul.
267. Anatidae indet. 9
Late Holocene – RS
Anatidae – Milheira et al. 2019: 45.
Milheira et al. ( 2019 ) reported anatid remains among the material from the sites PSG-02 and PSG-07 of the Pontal da Barra cerrito complex in Pelotas, Rio Grande do Sul.
Dendrocygninae Reichenbach
Dendrocygna Swainson
268. Dendrocygna sp.
Quaternary – MG
Dendrocycna sp. 1 v. 2 (vel D. viduata et sp. alia; vel sp. una, non D. viduata ) – Winge 1887: 19.
Dendrocygna viduata [in part] – Goeldi 1894: 603.
Dendrocygna spec. [in part] – Goeldi 1894: 603.
Dendrocygna sp. (“cfr. viduata und andere Arten”) – Lambrecht 1933: 740.
Dendrocygna viduata – Brodkorb 1964: 237.
Dendrocygna viduata – Cuello 1988: 34.
Dendrocygna sp. – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported the distal end of a tibiotarsus and a tarsometatarsus from Lapa da Escrivânia XI, the latter slightly thinner than in Dendrocygna viduata , and a humerus and a carpometacarpus from Lapa da Lagoa do Sumidouro, which correspond well to D. viduata . From the latter locality, there is also a femur that possibly fits here, but only bones of a mounted specimen (lacking the femora) were available for comparison. He noted it is not clear whether the bones from Lapa da Lagoa do Sumidouro are of D. viduata , and those from Lapa da Escrivânia XI are of a related species, or both materials belong to this related species.
Anatinae Leach
Neochen Oberholser
269. † Neochen pugil (Winge)
Quaternary – MG
Figs 19–21, 35A
Type locality: Depicted material from Lapa da Escrivânia V (Hansen 2012 ).
Etymology: pugil , Latin for “boxer” (Lambrecht 1933: 377).
“ Chauna eller Palamedea ” [in part] – Reinhardt 1881: 145.
“ Chauna or Palamedea ” [in part] – Reinhardt 1882: 325.
Chenalopex pugil n. sp., affinis jubatæ (Spix) sed multo major [new species] – Winge 1887: 19–22, figs 1–6.
Chenalopex pugil – Goeldi 1894: 603.
Alopochen pugil – Rothschild 1907: ix.
Chenalopex pugil [in part] – Lambrecht 1921: 11.
Chenalopex pugil [in part] – Lambrecht 1933: 376, 377, 390, 741, 880, 194A.
Neochen pugil – Howard 1964: 281–282, pl. VI: C, D [May 1964].
Neochen pugil – Brodkorb 1964: 216 [26 June 1964].
“Fighting shelduck, Neochen pugil ” – HRH The Prince Philip, Duke of Edinburgh and Fisher 1970: 193.
Neochen pugil [ Chenalopex pugil ] – Mones 1986: 81–82.
Neochen pugil – Cuello 1988: 10.
Neochen pugil – Alvarenga 1993a: 62.
Neochen pugil – Alvarenga 1997: 123.
“ Neochen pugil (Winge, 1887 ).—Greater Orinoco Sheldgoose” – Livezey 1997: 475.
Neochen pugil – Alvarenga 1998: 60.
Chenalopex pugil [original name] – Hansen 2012: 98–99, 41A, 41B.
Neochen pugil [in part] – Nascimento and Silveira 2020: 490, 3.
Neochen pugil – Agnolín et al. 2024: 249, 3H, F–G.
Winge ( 1887 ) described this species as Chenalopex pugil based on remains from three localities: several bones from Lapa da Escrivânia V, belonging to at least three males, more bones from Lapa da Escrivânia XI, mainly from a female, and some fragments associated with a male from Lapa dos Tatus. It constitutes the first species of fossil bird (and dinosaur) named for Brazil.
The material is represented by the cervical vertebrae, coracoid (not listed, but depicted and mentioned in the description), a rib, a fragment of the anterior part of the sternum, part of the synsacrum and pelvis, proximal and distal ends of the humerus, proximal end of the ulna, proximal and distal ends of the radius, carpometacarpus, first phalanx of the major digit of the wing, femur, distal end of the tibiotarsus, tarsometatarsus, and first phalanx of digit II and first phalanx of digit III of the foot. No elements were designed as a type (Howard 1964 ). However, a right coracoid (ZMUC 12115), the proximal end of a left humerus (ZMUC 12017), a right carpometacarpus (ZMUC 12044), a left tibiotarsus lacking the proximal end (ZMUC 12122), and a left tarsometatarsus (ZMUC 12084) associated with the male sex from Lapa da Escrivânia V were depicted. This material was listed as the type by Brodkorb ( 1964 ) and Hansen ( 2012 ).
Winge found it quite similar to Neochen jubata , but larger and different in some characters (e.g., in the cervical vertebrae, coracoid, sternum, humerus, and radius) and well-differentiated from Alopochen aegyptiaca , Plectropterus gambensis , Chloephaga melanoptera , and Chloephaga picta . There are two sets of bones, one of larger size and one smaller, which he believed belonged to males and females, respectively, and correspond almost precisely to the differences observed between two specimens of N. jubata that were available for comparison—a taxidermied specimen with no defined sex, but probably a male, and a disarticulated skeleton of a young female. The carpometacarpus and tarsometatarsus are proportionally longer, and the femur, pedal phalanges, and coracoid are proportionally slightly shorter than in N. jubata . Winge noted that, if both sexes of the two species could be appropriately compared, the differences observed in the radius and cervical vertebrae, and perhaps even in the sternum, could disappear. As in N. jubata , there is a well-developed spur in the alular metacarpal, probably used for fighting, and it likely inspired its specific name. Unregistered fragments in Lund’s collection belonging to this species were attributed to Chauna or Anhima by Reinhardt ( 1881 ), but no anhimid remains are known among the material from the Lagoa Santa region.
Winge proposed that this species might still be found alive in remote areas in the inner regions of Brazil, near the southern tributaries of the Amazon River. Nevertheless, so far there has been no record of its existence in historical times. Lambrecht ( 1933: 377) mistakenly commented that “Reinhardt collected the same from the Brazilian Campos”.
Andrews ( 1897: 350) made a parallel between the two living species of “ Chenalopex ”, C. aegyptiaca and C. jubata , where each is represented in its respective range by a much larger pleistocenic form, Centrornis majori and Chenalopex pugil , respectively. He noted they differ in the shape of the legs but have an almost identical wing structure, where the similarities between the carpometacarpi are a “remarkable example of parallel modification”, but not in a way that suggests a generic relationship between the two fossil taxa (Howard 1964 ).
Neochen pugil♂ , Lapa da Escrivânia V, holotype: proximal end of the left humerus (ZMUC 12017), cranial (A) and caudal (B) views; right coracoid (ZMUC 12115), ventral (C) and dorsal (D) views; right carpometacarpus (ZMUC 12044), ventral (E) and dorsal (F) views; left tibiotarsus lacking the proximal end (ZMUC 12122), cranial (G) and caudal (H) views; left tarsometatarsus (ZMUC 12084), dorsal (I) and plantar (J) views (K.L. Hansen). Scale bar: 10 mm.
Neochen pugil ♀, Lapa da Escrivânia XI: ulnae (A, B); carpometacarpus (C); femur (D); humeri (E, F, G). ♂, Lapa dos Tatus: coracoid (H), carpometacarpus (I), foot phalanx (J) (K.L. Hansen). Unspecified Lagoa Santa locality: right carpometacarpus (NHMUK PV OR 18906), proximal (K), distal (L), cranial (M), ventral (N), dorsal (O), and caudal (P) views (S. Chapman). Scale bar: 10 mm.
Neochen pugil ♂, Lapa da Escrivânia V: carpometacarpi (A, B, C, G, H); rib (D); partial synsacrum and pelvis (E); wing phalanges (I, J); vertebrae (F, K, L, O, P); coracoids (M, N, V); anterior part of the sternum (Q); ulnae (R, S); tarsometatarsi (T, Aa, Ab); pedal phalanx (U); radius (X); tibiotarsi (Y, Z, Ac); femora (W, Ad, Ae) (K.L. Hansen). Scale bar: 10 mm.
The first mentions of the combination Neochen pugil are those by Howard (May 1964 [29] ) and Brodkorb (26 June 1964 ). Howard noted that the “ Chenalopex ” species are recognized as Neochen in South America and Alopochen in Africa. Winge himself found the placement of Chenalopex jubata and C. pugil in the same genus as Chenalopex aegyptiaca debatable. Due to the differences in size and structure, Andrews ( 1897 ) suggested classifying C. pugil in a new genus apart from C. jubata . Rothschild ( 1907 ) listed it under Alopochen , possibly following the classification of C. jubata and C. aegyptiaca in this new genus by Stejneger ( 1885 ). The name Neochen was only proposed for the then Alopochen jubata by Oberholser in 1918 , without a diagnosis for the described fossil species of South America, Chenalopex pugil and the Argentinean Chenalopex debilis (now Neochen debilis ), of middle pleistocenic age (Agnolín 2006a ). A third fossil species, Neochen barbadiana , was described in 1965 from the late Pleistocene of Barbados (Brodkorb 1965 ). Agnolín et al. ( 2024 ) described the new species Chloephaga dabbenei (middle Pleistocene of Bajo San José, Buenos Aires Province, Argentina) and found similarities between it and N. pugil . They noted it is not improbable that the latter belongs to the genus Chloephaga .
Lydekker ( 1891 ) associated with this species a right carpometacarpus (NHMUK PV OR 18906) described as “imperfect” from the British Museum (Natural History) collection. The institution acquired this bone in 1848 as part of the Claussen Collection, and it comes from a cave in the same region as Lund’s finds. Lambrecht ( 1933 ) erroneously listed a tarsometatarsus present in that collection.
Cairina Fleming
270. Cairina moschata (Linnaeus)
Quaternary – MG, PI, RJ
Anas [in part] – Lund (in Winge 1887 ): 22.
Anatid. sp., s. under Cairina moschata [in part] – Winge 1887: 13.
Cairina moschata [in part] – Winge 1887: 14.
Cairina moschata (et forte sp. alia) – Winge 1887: 22.
Cairina moschata – Goeldi 1894: 603.
Cairina moschata – Lambrecht 1933: 743.
Cairina moschata – Brodkorb 1964: 238.
Cairina moschata – Cuello 1988: 35.
Cairina moschata – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported remains from three localities. From Lapa da Pedra dos Índios, a humerus (“ Anas ” in Lund’s catalog). From Lapa da Lagoa do Sumidouro, an ulna and a tibiotarsus, besides the first part of pedal digit III that possibly belongs to this species and the distal end of a tibiotarsus and the proximal end of a carpometacarpus that possibly belong to a female. Finally, from Lapa da Escrivânia XI, several bones, including a humerus he considered with much uncertainty that might belong to a small female.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Kneip et al. ( 1994 ) reported a bone from layer I (younger than 1,790±50 years BP) and another from layer III (1,810±40 years BP) of Sambaqui da Pontinha in Saquarema, Rio de Janeiro.
Amazonetta Boetticher
271. Amazonetta brasiliensis (Gmelin)
Quaternary – MG, PI, RJ
Anas sp., s. Anm. under 14. Anas brasiliensis [in part] – Winge 1887: 15.
Anas brasiliensis (et aff.) – Winge 1887: 22.
Querquedula brasiliensis – Goeldi 1894: 603.
Anas brasiliensis – Lambrecht 1933: 740.
Nettion brasiliense – Brodkorb 1964: 243.
Nettion brasiliense [ Anas brasiliensis ] – Mones 1986: 81–82.
Amazonetta brasiliensis – Cuello 1988: 35.
Amazonetta brasiliensis – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, including an uncertain femur, several bones likely of a single individual from Lapa da Escrivânia XI, and some bones from Lapa da Lagoa do Sumidouro. The material is represented by the coracoid, humerus, ulna, radius, carpometacarpus, femur, tibiotarsus, and tarsometatarsus. Additionally, from Lapa da Escrivânia V, there is the distal end of a humerus that is quite similar to that of Amazonetta brasiliensis but larger, besides a coracoid and a carpometacarpus that perhaps can also be assigned here. From “various caves” [26] , two coracoids, slightly larger than in A. brasiliensis and different from it and each other (even the genus assignment is uncertain), besides a cervical vertebra of a considerably large anatid.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Kneip et al. ( 1994 ) reported a bone from layer I (3,610±190 years BP) of Sambaqui do Moa, and two bones of at least one individual from layer I (younger than 1,790±50 years BP) and two bones of at least one individual from layer III (1,810±40 years BP) of Sambaqui da Pontinha in Saquarema, Rio de Janeiro.
Netta Kaup
272. cf. Netta peposaca (Vieillot)
Late Holocene – SC
“marrecão” – Schmitz and Verardi 1996: 143.
Schmitz and Verardi ( 1996 ) reported “marrecão” (= Netta peposaca ?) remains from the Cabeçudas site in Itajaí, Santa Catarina.
273. Netta peposaca (Vieillot)
Rosa (in Schmitz et al. 1997 ) reported a single fragment with burn traces from the RS-170 site in Santa Vitória do Palmar, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Mergus Linnaeus
274. Mergus cf. octosetaceus Vieillot
Quaternary – MG
Mergus sp., verisimillime M. octosetaceus Vieill. ( brasilianus Vieill. postea) – Winge 1887: 23.
Mergus spec. ( octosetaceus ?) – Goeldi 1894: 603.
Mergus sp. (“cfr. M. octosetaceus ”) – Lambrecht 1933: 736.
Mergus octosetaceus – Brodkorb 1964: 249.
Mergus octosetaceus – Mones 1986: 82.
Mergus octosetaceus – Cuello 1988: 36.
Mergus cf. octosetaceus – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported two humeri and two tarsometatarsi from Lapa da Escrivânia V, besides the distal end of a tibiotarsus that probably belongs to the same species, possibly Mergus octosetaceus , of which there were no records in Minas Gerais. He noted that the tarsometatarsus is similar to that of Mergus merganser , Mergus serrator , and Mergellus albellus , shorter and stouter than in Merganetta armata , and longer than in Lophodytes cucullatus . The humerus has almost the same size as in M. albellus (but the tarsometatarsus is larger) and is slightly larger than in Oxyura leucocephala (of which the humerus used in comparison lacked the proximal end).
Nomonyx Ridgway
275. Nomonyx dominicus (Linnaeus)
Quaternary – MG
Erismatura dominica – Winge 1887: 22.
Erismatura dominica – Goeldi 1894: 603.
Erismatura dominica – Lambrecht 1933: 736.
Nomonyx dominicus – Brodkorb 1964: 248.
Nomonyx dominicus [ Erismatura dominica ] – Mones 1986: 81–82.
Oxyura dominica – Cuello 1988: 36.
Nomonyx dominicus – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported a tarsometatarsus from Lapa da Escrivânia V, which is very similar to the one of a recent specimen, and a femur of a small duck from Lapa da Lagoa do Sumidouro, which possibly belongs to this species.
Galliformes Linnaeus
276. Galliformes indet.
Early Holocene – SP
“galeiformes” [sic] – Alves 2008: 170.
Alves ( 2008 ) reported galliform remains (at least one individual) from area III of the Capelinha I site in Cajati, São Paulo, aged 9,250±50 years BP.
Cracidae Rafinesque
277. cf. Cracidae
Rosa ( 2009 ) reported a possible cracid bone from square A6 of the third occupation of the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. The material is deposited in the CEPA collection.
278. Cracidae indet.
Holocene – RS
Cracidae – Jacobus 2004: 99.
Jacobus ( 2004 ) reported cracid remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
Penelope Merrem
279. Penelope sp. 1
Quaternary – MG
Penelope sp. (fere certe sp. 2, vel 3) [in part] – Winge 1887: 23–24.
“duas ou tres especies de Penelope ” [in part] – Goeldi 1894: 440.
Penelope sp. (“cfr. superciliosus Ill. oder vetula ”) [in part] – Lambrecht 1933: 752.
Penelope obscura – Brodkorb 1964: 306.
Penelope obscura – Mones 1986: 85.
Penelope obscura – Cuello 1988: 40.
Penelope sp. 1 – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) divided the material referrable to the genus Penelope into three sets, based on the size of the bones, which can represent up to three species. He had available for comparison Penelope superciliaris and a larger species, “ Penelope cristata ?”, which he believed to be well determined. The first and larger set consists of two right femora, much larger than in P. superciliaris and similar to “ P. cristata ”, from “various caves” [26] , and the lower end of a coracoid, of equivalent size to the femora, from “a saltpeter cave near Escrivânia”—erroneously listed as “Lapa da Escrivânia” by Brodkorb ( 1964 ) and Cuello ( 1988 ).
280. Penelope sp. 2
Quaternary – MG
Penelope [in part] – Lund (in Winge 1887 ): 23.
Penelope sp. (fere certe sp. 2, vel 3) [in part] – Winge 1887: 23–24.
“duas ou tres especies de Penelope ” [in part] – Goeldi 1894: 440.
Penelope sp. (“cfr. superciliosus Ill. oder vetula ”) [in part] – Lambrecht 1933: 752.
Penelope superciliaris – Brodkorb 1964: 306.
Penelope superciliaris – Mones 1986: 85.
Penelope superciliaris – Cuello 1988: 40.
Penelope sp. 2 – Nascimento and Silveira 2020: 490.
The set of medium-sized bones attributed to Penelope by Winge ( 1887 ) consists of material from two sites. From Lapa do Baú, two left humeri (“ Penelope ” in Lund’s catalog)—one without recorded origin but with a very similar aspect to the other, both being larger than in P. superciliaris and slightly smaller than in “ P. cristata ” and differ slightly from both. From Lapa da Escrivânia III, bones of a single individual: mandible, a cervical vertebra, humerus, ulna, first phalanx of the major digit of the wing, femur, tibiotarsus, tarsometatarsus, and some pedal phalanges. They are slightly larger than P. superciliaris and slightly smaller than “ P. cristata ”. The humerus is very similar to that from Lapa do Baú. The tibiotarsus is slightly thinner and somewhat shorter than in “ P. cristata ”, and the first phalanx of the major digit of the wing and the pedal phalanges are as large as in it.
281. Penelope sp. 3
Quaternary – MG
Falco [in part] – Lund (in Winge 1887 ): 24.
Penelope sp. (fere certe sp. 2, vel 3) [in part] – Winge 1887: 23–24.
“duas ou tres especies de Penelope ” [in part] – Goeldi 1894: 440.
Penelope sp. (“cfr. superciliosus Ill. oder vetula ”) [in part] – Lambrecht 1933: 752.
Ortalis guttata – Brodkorb 1964: 306.
Ortalis guttata – Mones 1986: 85.
Ortalis guttata – Cuello 1988: 40.
Penelope sp. 3 – Nascimento and Silveira 2020: 490.
The third, smaller-sized set of bones attributed to Penelope by Winge ( 1887 ) consists of remains from three localities. From Lapa da Escrivânia V, the distal end of a humerus, slightly smaller than in Penelope superciliaris , but probably of this species. From “various caves” [26] , two coracoids of young individuals, one of which is very similar to P. superciliaris but somewhat larger and the other is shorter and slightly different. Finally, from a “cave near Mocambo”, a humerus lacking the proximal end (“ Falco ” in Lund’s catalog), slightly larger and divergent when compared with P. superciliaris .
Brodkorb ( 1964 ), probably based on the sizes, associated the three sets, from the largest to the smallest, respectively, with Penelope obscura , P. superciliaris , and Ortalis guttata .
282. Penelope sp. 4
Guérin et al. ( 1996 ) reported material attributable to Penelope superciliaris or Penelope jacucaca from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
283. Penelope sp. 5
Jacobus and Rosa ( 2013 ) reported Penelope remains from the Sangão (RS-S-327) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul. Previously, this record was briefly mentioned by Dias ( 2003 , 2004a ).
284. † Penelope spp. 6
Alvarenga and Silveira (in Silveira et al. 2008 ) reported undescribed species of Penelope among the approximately 20,000-year-old fossils found in caves of Minas Gerais and Bahia.
285. Penelope sp. 7
Jacobus and Rosa ( 2013 ) reported Penelope remains from the Deobaldino (RS-S-395) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
286. Penelope sp. 8
Holocene – RS
Penelope sp. – Jacobus and Rosa 2013: 247.
Jacobus and Rosa ( 2013 ) reported Penelope remains from the Schneider (RS-C-14) rock shelter in São Sebastião do Caí, Rio Grande do Sul.
287. Penelope sp. 9
Holocene – RS
Penelope sp. – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported Penelope remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
288. Penelope sp. 10
Jacobus and Rosa ( 2013 ) reported Penelope remains from the Garivaldino (RS-TQ-58) rock shelter in Montenegro, Rio Grande do Sul. The material is deposited in the CEPA collection.
289. Penelope sp. 11
Middle Holocene – SC
Penelope sp. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) associated with this genus a bone from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
290. Penelope superciliaris Temminck
Quaternary – MG
Penelope superciliaris sp. – Sales 2003: 214.
Sales ( 2003 ) reported two specimens among the bones and mummified carcasses found between 2001 and 2002 in Zona II of Lapa do Rezar in Itacarambi, Minas Gerais. Regarding the total material, he specified that 18 bone sets and five mummified carcasses were found but did not attribute the material’s nature to each species or age.
291. cf. Penelope obscura Temminck
Beber ( 2004 , 2005 ) reported avian remains that possibly belong to Penelope obscura from the Perau das Cabeças (RS-A-28) rock shelter in Vacaria, Rio Grande do Sul.
292. Penelope jacucaca Spix
Holocene – PI
Penelope jacucaca – Barbosa 2017: 125.
Barbosa ( 2017 ) reported this species from Toca do Alto da Serra do Capim in Guaribas, Piauí. Seven bones were found in association with combustion structures.
Crax Linnaeus
293. Crax sp.
Quaternary – MG
Crax ? [in part] – Lund (in Winge 1887 ): 24.
Crax sp. 1 vel 2 – Winge 1887: 24.
“uma ou duas [especies] de Crax ” – Goeldi 1894: 440.
Crax sp. – Lambrecht 1933: 752.
Crax fasciolata – Brodkorb 1964: 306.
Crax fasciolata – Mones 1986: 85.
Crax fasciolata – Cuello 1988: 40–41.
Crax sp. – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) described material he considered to be one or two Crax species. From Lapa da Escrivânia V, a coracoid, the proximal and distal ends of a tibiotarsus, and part of a tarsometatarsus, all larger than in a recent skeleton. From Lapa da Escrivânia XI, part of the proximal end of a humerus and a tibiotarsus (approximately the same size as the one from Lapa da Escrivânia V). From Lapa do Capão Seco, a humerus (“ Crax ?” in Lund’s catalog) and a tibiotarsus, of similar size to that observed in recent skeletons (although the humerus is more robust) but considerably smaller than the material of the other caves. Finally, from “various caves” [26] , the distal end of an ulna, slightly larger than in recent skeletons. Only two recent skeletons with a similar size were available for comparison but without a reliable identification of the species.
Odontophoridae Gould
Odontophorus Vieillot
294. Odontophorus capueira (Spix)
Quaternary – MG, RS
Perdix dentata [in part] – Lund 1841d: 18.
Perdix [?; in part] – Giebel 1846: 312.
“Überreste einer der P. dentata nah verwandten Art” [in part] – Giebel 1847: 23.
Perdix [in part] – Pictet 1853: 415.
“les Perdix” [in part] – Liais 1872: 303.
Perdix aff. dentatæ [in part] – Lund (in Winge 1887 ): 24.
Odontophorus dentatus – Winge 1887: 24–25.
Odontophorus dentatus – Goeldi 1894: 440.
Odontophorus dentatus – Lambrecht 1933: 752.
Odontophorus gujanensis – Brodkorb 1964: 327.
Odontophorus gujanensis [ Odontophorus dentatus ] – Mones 1986: 85–86.
Odontophorus gujanensis – Cuello 1988: 41.
Odontophorus gujanensis (= O. dentatus ) – Tambussi and Noriega 1996: 251.
Odontophorus capueira – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported remains from various sites. From Lapa da Escrivânia V, a tarsometatarsus. From Lapa do Marinho II, two paired humeri and two tibiotarsi. From Lapa do Periperi I, a coracoid and an ulna. From Lapa Vermelha (Lagoa Santa), a humerus (“ Perdix aff. dentatæ ” in Lund’s catalog). From “various caves” [26] , a tibiotarsus and a tarsometatarsus. Finally, of unknown origin, several humeri (two of them determined as “ Perdix aff. dentatæ ” from “caves in Mocambo” in Lund’s catalog), a coracoid, and a tarsometatarsus. He further noted that some of these humeri are probably from Lapa da Escrivânia V.
This material was listed as Odontophorus gujanensis by Brodkorb ( 1964 ), which was followed by Mones ( 1986 ) and Cuello ( 1988 ), but this species is not found, at least in the present, in Minas Gerais. Winge probably based the name Odontophorus dentatus (Temm.) on Reinhardt’s ( 1870 ) Odontophorus dentatus (Licht.). Krabbe ( 2007 ) reviewed the skins collected by Lund and Reinhardt and associated the name used by Reinhardt with Odontophorus capueira capueira .
Jacobus ( 2004 ) reported remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
†Quercymegapodiidae Mourer-Chauviré
† Taubacrex Alvarenga
295. † Taubacrex granivora Alvarenga
Late Oligocene/Early Miocene – SP
Fig. 33 A
Type locality: Tremembé Formation, Taubaté basin, municipality of Tremembé, State of São Paulo, Brazil.
Etymology: Taubacrex , a portmanteau of Taubaté and the Greek krex (a rallid), and granivora , Latin for “grain eater”.
Rallidae – Castro et al. 1988b: 2362.
Taubacrex granívora sp. n. [new genus and species] – Alvarenga 1988: 321–328, figs 1–3, 6.
Taubacrex granivora – Alvarenga 1993a: 63.
Taubacrex granivora – Alvarenga 1997: 123.
Taubracrex granivora – Kellner 1998: 654, 659.
Taubacrex granivora – Alvarenga and Höfling 2000: 589.
Taubacrex granivora – Alvarenga and Höfling 2004: 827.
Ameripodius granivora – Mourer-Chauviré et al. 2011: 620.
Taubacrex granivora – Alvarenga and Höfling 2011: 128.
Taubacrex granivora – Carmo et al. 2024: 5.
Alvarenga ( 1988 ) described this new genus and species from remains preserved in a pyrobituminous shale fragment of the Tremembé Formation. It was collected at a depth of about 6 m at the Mina Nossa Senhora da Guia by Petrobrás engineer Mustafá Hanzagic in 1950. Its appearance indicates the remains were preserved completely, but parts of the skeleton were lost, possibly during their collection.
The specimen consists of an incomplete and fragmented post-cranial skeleton (IG-208-V, holotype) with bone remains and impressions including the right coracoid, a humerus, the left radius, the left pelvis, the femora, the tibiotarsi, and the left tarsometatarsus, aside from ossified tendons in the tibiotarsus and tarsometatarsus. The preservation of the gizzard with nine spheroidal gastroliths and seed impressions indicates its eating habits and consists of the oldest record of this nature for the Galliformes (Mayr 2009 ). There are also feather impressions with a carbonized/imprint preservation (Prado et al. 2016a ) without relief, and they appear to be small tectrices and upper wing coverts, and some larger ones may be secondary remiges. Primary remiges or larger rectrices are absent. A preserved ungual phalanx and the impression of another with relatively short and slightly curved claws suggest, along with other features of the skeleton, that it was cursorial, although capable of flying. Its dimensions were close to those of Gallinula galeata . The fragment of the shale containing the fossil has a triangular shape and measures about 22 x 20 cm. It was originally published as part of the Seção de Paleontologia e Estratigrafia of IG (now Museu Geológico de São Paulo, Instituto de Pesquisas Ambientais), but it could not be found in that collection nor its registers as of 2023 (IPA – Núcleo de Museus, Acervos Arquivísticos e Iconográfico, personal communication) and its whereabouts are unknown.
Alvarenga placed the taxon within Rallidae (Gruiformes) and pointed out that similarities with Galliformes were observed but always secondarily to the previous group. Mourer-Chauviré ( 2000 ) reanalyzed the material (including comparison with Ameripodius from France) and attributed it to Quercymegapodiidae. Contemporarily, Alvarenga and Höfling ( 2000 ) also suggested a classification as a galliform closer to the megapodiids. It was mentioned as Ameripodius granivora by Mourer-Chauviré et al. ( 2011 ) and was followed by Tambussi and Degrange ( 2013 ). This combination, however, was possibly a mistake (C. Mourer-Chauviré, personal communication). Mayr ( 2009 , 2022 ) noted that even though T. granivora and Ameripodius silvasantosi differ in the shape of the coracoid (Mourer-Chauviré 2000 ), this bone is slightly crushed in the holotype of the former, and a direct comparison of the actual specimens is necessary to exclude the possibility that the two are conspecific.
† Ameripodius Alvarenga
296. † Ameripodius silvasantosi Alvarenga
Late Oligocene/Early Miocene – SP
Fig. 33 B
Type locality: State of São Paulo, municipality of Tremembé, Santa Fé Farm; Taubaté Basin, Tremembé Formation, uppermost layer of pyrobituminous shales at indeterminate level.
Etymology: Ameripodius , a portmanteau of America and the Greek podius (foot), referring to the similarity to the Megapodiidae, and silvasantosi , honoring Brazilian paleoichthyologist Rubens da Silva Santos (1918–1996).
Galliformes – Alvarenga (in Mourer-Chauviré 1992b ): 7, 8.
Galliformes – Alvarenga 1993b: 24, 18.
Ameripodius silvasantosi sp. n. [new genus and species] – Alvarenga 1995b: 33–44, figs 2–8.
Ameripodius silvasantosi – Alvarenga 1997: 123.
Ameripodius silvasantosi – Kellner 1998: 654, 659.
Ameripodius silvasantosi – Alvarenga and Höfling 2000: 589.
Ameripodius silvasantosi – Alvarenga and Höfling 2004: 827.
Ameripodius silvasantosi – Alvarenga and Höfling 2011: 128.
Ameripodius silvasantosi – Carmo et al. 2024: 5.
Alvarenga ( 1995b ) described this new genus and species from the partial impression of a skeleton found in the Tremembé Formation of the Fazenda Santa Fé. The fossil was found in July 1989 through the cooperative work of Alvarenga, Rubens da Silva Santos, and other professors from UERJ. It was then considered the first record of a galliform for the Tertiary of South America (Alvarenga 1993b , 1995b ; see Taubacrex ).
The material (MN 4488-V, holotype) consists of part of the shoulder girdle and wings, preserved in a pyrobituminous shale plate divided into part and counterpart. The cranial half of the right coracoid, the left coracoid, and the left carpometacarpus were almost completely removed from the shale. The humeri were removed in fragments, leaving an impression cast. Partial impressions of the right radius, ulna, and carpometacarpus were also obtained. The anterior margin of the sternal keel, the left clavicle including the symphysis region, the fragmented and incomplete right scapula, the almost complete left radius, and the phalanges 1 and 2 of the major digit of the left wing were embedded in the matrix. The relatively robust and slightly arched ulna suggests its wings were proportionally short and strong. Its overall dimensions were that of an Odontophorus -sized bird.
Alvarenga attributed it to the Quercymegapodiidae, which at the time included only Quercymegapodius depereti from the late Eocene and Quercymegapodius brodkorbi from the middle and late Eocene, both from Phosphorites du Quercy, France (Mourer-Chauviré 1992a , 2006 ). A second, larger species of the Brazilian genus was described by Mourer-Chauviré ( 2000 ) as Ameripodius alexis , from the early Miocene of Saint-Gérand-le-Puy, France. The presence of this genus in both continents emphasizes the similarity between the early Paleogene South American and European avifaunas (Mourer-Chauviré 2000 ). Mourer-Chauviré ( 1999 , 2000 ) hypothesized that, like the Megapodiidae, which are capable of crossing large extensions of water and colonizing islands, there was a possible direct dispersion for Ameripodius across the ocean between Europe and South America, or vice versa, when the Atlantic was considerably narrower than it is today. According to Mayr ( 2016 ), the very narrow carpometacarpus of the quercymegapodiids indicates they differed from the living Galliformes in their flight capacity, which agrees with the fact that they are the only stem group of the clade known to have reached the geographically isolated South American continent.
†Odontopterygiformes Howard
†Pelagornithidae Fürbringer
† Pelagornis Lartet
297. † Pelagornis longirostris (Spulski)
Paleogene? – BR?
Type locality: None attributed.
Etymology: Pelagornis , Greek for “sea bird”, and longirostris , Latin for “long face”.
Odontopteryx longirostris [new species] – Spulski 1910: 507–521, figs 1–7.
Odontopteryx longirostris – Lambrecht 1921: 41.
Pseudodontornis longirostris – Lambrecht 1930: 1–17, figs 3, 6, pls. 1–2.
Pseudodontornis longirostris – Lambrecht 1933: 305–307, figs 110–111.
Pseudodontornis longirostris – Brodkorb 1963: 263.
Pseudodontornis longirostris – Fisher 1967: 739.
Pseudodontornis longirostris – Harrison and Walker 1976: 15–17, figs 15C, 17D, 21A, 24, 25B, 25D, 26B, 30B.
Pseudodontornis longirostris [“ Odontopteryx ” longirostris ] – Mones 1986: 79.
Pseudodontornislongirostris – Bocheński 1997: 300.
Pelagornislongirostris – Mayr and Rubilar-Rogers 2010: 1327.
Pelagornis ( Pseudodontornis ) longirostris – Ksepka 2014: 10624.
Pseudodontornis longirostris – Mayr 2022: 44, 4.2A.
Spulski ( 1910 ) erected the species Odontopteryx longirostris for an incomplete, 40-cm skull and jaw of unknown age and origin, characterized by pseudoteeth. The fossil also had conserved the right eye’s sclerotic ring, which was lost in an accident before the analysis. The specimen was supposedly brought from Brazil to Germany by a sailor five years earlier, who sold it to the rarities dealer J. Schulze in Königsberg, and it was then acquired in 1905 by Prof. Braun for the Zoologische Institut.
Spulski compared it with Odontopteryx toliapica from the early Eocene of London Clay described by Richard Owen in 1873. However, after further preparation and study of the specimen, in addition to also examining Owen’s fossil, Lambrecht ( 1930 ) found it distinct enough to be classified into the new genus Pseudodontornis , based on the pseudoteeth and other characteristics.
Spulski noted the orbits were filled with pure and granular limestone and speculated the fossil could date from the Eocene, based on the significant similarity he found with the Odontopteryx toliapica skull, as well as its “primitive characteristics”. Lambrecht disagreed and mentioned that the mere presence of pseudoteeth is not a sufficient argument and the Pseudodontornis longirostris skull is quite specialized, and these two isolated points cannot serve as a basis to determine its age.
During the fossil reanalysis, Lambrecht presented a portion of the matrix to Prof. Friedrich von Huene, who had returned from South America, and asked his opinion on its origin. He was, however, unaware of any similar rock. Lambrecht achieved the same result by showing Prof. von Fryberg, who had been to Brazil three times. Due to the “impression of being surprisingly new” and the highly crystalline character of the rock, Lambrecht discarded an origin associated with freshwater. The presence of brachiopod traces reinforced the association with a marine coastal environment. On its probable age, he also observed a negative impression of a bivalve shell, “close to Cardium ( Laevicardium ) cingulatum ”, from the late Oligocene of Europe (“but that also occurred in the Pliocene”) but noted that it might be a secondary deposition in the matrix.
Lambrecht ( 1930 , 1933 ) did not exclude the possibility that the fossil was found in Germany and was attributed to Brazil so that it could be sold at a higher price. However, the pelagornithids had a worldwide distribution from the late Paleocene to the late Pliocene (Mayr 2016 ), including records from four Neogene formations in South America—in Chile, Peru, and Venezuela (Solórzano and Rincón 2015 )—and it would not be unlikely that representatives of the group had inhabited Brazil, especially given its extensive coastline. Mayr and Rubilar-Rogers ( 2010 ) speculated that the species probably lived during the Neogene. Brodkorb ( 1963 ) suggested a Miocene age and German origin. Mones ( 1986 ) associated it, with uncertainty, with the Eocene. Mlíkovský ( 2002 ), in contrast to Brodkorb’s assumption, argued there are no suitable locations of Miocene age in that part of Europe, and that if the fossil indeed came from that continent, it would have been from the Eocene marine deposits in the North Sea Basin, which are well represented in northern Germany, Denmark, and England.
Mayr ( 2022 ) argued against a Brazilian origin for the fossil. His assumption was based on the fact that it was initially described in the early twentieth century and is preserved in what seems to be a fairly solid matrix, which indicates it may come from the London Clay of Isle of Sheppey in southern England. If this is the case, it would possibly belong to Dasornis emuinus , of similar size and morphology.
The specimen was deposited in the Geological-Paleontological Museum of the Institutes der Albertus-Universität Königsberg, then in Prussia. Königsberg was heavily bombed by the Allies during World War II and, by the end of the war, it was annexed to the Soviet Union and renamed Kaliningrad. The fossil appears to have been destroyed during these events (Mayr and Rubilar-Rogers 2010 , Ksepka and Habib 2016 ). Harrison and Walker ( 1976 ) tried to locate the specimen with the help of Dr. E.N. Kurotchkin (Paleontological Museum of the Moscow Academy of Sciences) but had no success.
Pelagornithid remains were reported from South Carolina in the USA by Hopson ( 1964 ), including a fragment of the mandible which he associated with Pseudodontornis longirostris . It was initially attributed to the early Miocene but probably comes from the late Oligocene according to Olson ( 1985b ). Hopson commented that the discovery of this material strengthens the assumption that the type specimen came from the Western Hemisphere, but not necessarily from North America, and argued that an oceanic bird of this size probably had a wide distribution. In addition, the distal portion of a tarsometatarsus he attributed almost certainly to Palaeochenoides mioceanus —a species initially described from an incomplete femur from the same deposit by Shufeldt in 1916 —was later considered to belong to P. longirostris by Howard and Warter ( 1969 ) and Harrison and Walker ( 1976 ).
Pelagornis longirostris: incomplete skull in lateral (A), ventral (B), and caudal (C) views as reprepared by Lambrecht; skull as prepared by Spulski (D) (Therese Domok, modified from Lambrecht 1930). Scale bar for A-C: 50 mm.
Gamble ( 1985 ) associated a jaw fragment from the early Paleogene Oldhaven Formation in Shelford Sandpit, England, with P. longirostris . The specimen, as well as another indeterminate jaw fragment associated with the Odontopterygiformes, does not appear to be of a bird but are possibly fish remains according to Harrison ( 1985 ).
Chávez and Stucchi ( 2002 ) associated cranial remains from the Bahía Inglesa Formation, northern Chile, with Pseudodontornis cf. longirostris . However, the same fossils, dating from the middle Miocene, have been reassigned to Pelagornithidae indet. cf. Pelagornis by Chávez et al. ( 2007 ), who adopted more conservative criteria while waiting for new materials.
Further materials were associated with Pseudodontornis , but their classification is uncertain due to the generally poor preservation state of this group’s fossils (Mayr and Rubilar-Rogers 2010 ). In New Zealand, remains found on the South Island were originally described as Pseudodontornis stirtoni by Howard and Warter ( 1969 ), of Miocene or Pliocene age (Howard and Warter 1969 , McKee 1985 ). From North Island, McKee ( 1985 ) reported unidentified pelagornithid remains from the middle Pliocene, which he thought might belong to Pseudodontornis , as it was the only genus described for the country. In England, Pseudodontornis tenuirostris was described from the late Paleocene by Harrison ( 1985 ). From Kazakhstan, of the same age, Pseudodontornis tshulensis was described by Averianov et al. ( 1991 ), both forms with questionable validity due to little knowledge about the intraspecific variability of pseudoteeth in pelagornithids (Mayr 2009 ). From the early Eocene of England, there is also Pseudodontornis longidendata described by Harrison and Walker ( 1976 ), which is possibly synonymous with Dasornis emuinus (Bowerbank 1854) according to Mayr ( 2008 , 2009 ). The genus was also mistakenly listed (in place of “pseudodontornids”) by Becker ( 1987 apud Goedert 1989 ) for the Miocene of Astoria Formation, central coast of Oregon, USA.
Olson ( 1985b ) noted the possibility that Pseudodontornis is synonymous with Pelagornis , a genus originally proposed by Lartet in 1857 to Pelagornis miocaenus from the Miocene of France. Mayr and Rubilar-Rogers ( 2010 ), by reviewing the taxonomy of the Neogene pelagornithids, considering the present knowledge, and aiming at clarity in the study of the materials, proposed that all its components be classified under Pelagornis . This would include, in addition to P. miocaenus , the following species then recognized as valid: P. longirostris , P. stirtoni , P. orri from the Miocene of California, P. mauretanicus from the Pliocene of Africa, and P. chilensis from the Miocene of Chile (see also Mayr et al. 2013 ). Pelagornis sandersi , notable for its enormous wingspan, was later described from material from the Oligocene of South Carolina (Ksepka 2014 ).
Phoenicopteriformes Fürbringer
Phoenicopteridae Bonaparte
† Agnopterus Milne-Edwards
298. † Agnopterus sicki Alvarenga
Late Oligocene/Early Miocene – SP
Fig. 33 C
Type locality: Brazil, state of São Paulo, municipality of Tremembé, Santa Fé Farm, Taubaté Basin, Tremembé Formation; layer of montmorillonite clay below the uppermost strata of pyrobituminous shales.
Etymology: Agnopterus , Greek for “unknown wing” (Milne-Edwards 1869 –1871: 84), and sicki , honoring German-Brazilian ornithologist Helmut Sick (1910–1991).
Agnopterus sicki sp.n. [new species] – Alvarenga 1990: 335–345, figs 1–2.
Agnopterus sicki Alvarenga 1990 – Alvarenga 1997: 123.
Agnopterus sicki – Kellner 1998: 654, 659.
Agnopterus – Alvarenga and Höfling 2000: 591.
Agnopterus – Alvarenga and Höfling 2004: 829.
Agnopterus sicki – Alvarenga and Höfling 2011: 127.
Agnopterus sicki – Carmo et al. 2024: 5.
Alvarenga ( 1990 ) described this new species from remains collected between 1984 and 1989 in the montmorillonite clays below the uppermost pyrobituminous shales of the Tremembé Formation in the Fazenda Santa Fé.
The material consists of a well-preserved distal fragment of the right tibiotarsus (holotype, MN 4257-V), possibly from an individual that had not reached adulthood, and a diaphysis fragment (MN 4258-V), which perhaps belongs to the same bone.
The genus Agnopterus was erected by Milne-Edwards ( 1869 –1871) for Agnopterus laurillardi from the late Eocene of France. Tugarinov ( 1940 ) described a second species, Agnopterus turgaiensis , from the late Oligocene of Kazakhstan. The Brazilian species is notably similar to the latter in morphology and size. Alvarenga noted that the genus must be revised. The inclusion of A. turgaiensis was questioned by Olson and Feduccia ( 1980 ) due to the significant age difference from the material described by Milne-Edwards. Mayr ( 2009 , 2022 ) also commented that A. sicki and A. turgaiensis are based on too fragmentary material for a reliable taxonomic assignment within the Phoenicopteriformes. Another species, Agnopterus hantoniensis , was associated with this genus by Lydekker ( 1891 ) from material of the late Eocene of England, but its status is uncertain (Mayr 2009 , Zelenkov 2013 ). Mayr ( 2009 ) considered it a stem member of the Phoenicopteriformes.
†Palaelodidae Stejneger
† Palaelodus Milne-Edwards
299. † Palaelodus aff. ambiguus Milne-Edwards
Late Oligocene/Early Miocene – SP
Fig. 33 D
Palaelodus aff. ambiguus – Alvarenga 1990: 335–345, figs 3–9.
Palaelodus – Alvarenga 1993a: 63.
Palaelodus sp. – Alvarenga 1993b: 24, 17.
Palaelodus – Alvarenga (in Sick 1993 ): 152.
Palaelodus sp. – Alvarenga 1997: 123.
Palaelodus – Alvarenga (in Sick 1997 ): 226.
Palaelodus aff. ambiguus – Kellner 1998: 654, 659.
Palaelodus – Alvarenga and Höfling 2000: 591.
Palaelodus – Alvarenga and Höfling 2004: 829.
Palaelodus – Alvarenga and Höfling 2011: 129.
Palaelodus aff. ambiguous [lapsus] – Tambussi and Degrange 2013: 44.
Palaelodus aff. P. ambiguus – Carmo et al. 2024: 5.
Besides the material described as Agnopterus sicki , Alvarenga ( 1990 ) associated five other dissociated fragments with the genus Palaelodus , which show great affinity in size and form with Palaelodus ambiguus . They may represent a new species, but the poor state of preservation prevents a more specific diagnosis.
The material consists of the distal end of a left tibiotarsus (MN 4259-V), the distal end of a right tarsometatarsus lacking the lateral trochlea (MN 4261-V), a right coracoid lacking the upper end and the lateral half of the sternum joint (MN 4262-V), the proximal end of a left tibiotarsus (MN 4260-V), and the proximal half of a left first phalanx of the wing (MN 4263-V).
The genus Palaelodus is represented by several species: Palaelodus ambiguus , Palaelodus crassipes , and Palaelodus gracilipes in Europe, described initially from the early Miocene of France, with the validity of the last two questioned; Palaelodus aotearoa from the early Miocene of New Zealand; Palaelodus kurochkini from the middle Miocene of Mongolia; and Palaelodus pledgei and Palaelodus wilsoni from the late Oligocene to the middle Miocene of Australia, where a specimen from the middle Pleistocene was also associated with P. wilsoni , but its specific status is still uncertain (Baird and Vickers-Rich 1998 , Worthy et al. 2010 , Zelenkov 2013 ). In the late Oligocene or early Miocene, the palaelodids had already reached an almost global distribution (Mayr 2016 ).
Noriega and Agnolín ( 2008 ) reported the distal end of a right tarsometatarsus (MACN PV 12756) from the “Mesopotamiense”, Ituzangó Formation (late Miocene) of Entre Ríos Province, Argentina. It was referred to as Palaelodus cf. ambiguus , and its measurements and morphology coincide with the Brazilian material.
Podicipediformes Fürbringer
Podicipedidae Bonaparte
300. †Podicipedidae indet. 1
Alvarenga ( 1993b , 1997 ) mentioned poorly preserved, undescribed podiciped remains from the Tremembé Formation that await the discovery of new specimens for further study. Alvarenga (in Castro et al. 1988b ) mentioned indeterminate anatid remains that appear to be reassigned to podicipedids in subsequent mentions.
301. Podicipedidae indet. 2
Carvalho ( 1984 ) reported podicipedid remains among the material discovered in 1978 at the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
302. Podicipedidae indet. 3
Late Holocene – RS
Podicipedidae – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported this family from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Tachybaptus Reichenbach
303. Tachybaptus dominicus (Linnaeus)
Quaternary – MG
Tachybaptes dominicus – Winge 1887: 25.
Podiceps ( Tachybaptes ) dominicus – Goeldi 1894: 603.
Tachybaptes ( Podicipes ) dominicus – Lambrecht 1933: 731.
Podiceps dominicus – Brodkorb 1963: 228.
Podicipes dominicus [ Tachybaptes dominicus ] – Mones 1986: 76.
Tachybaptus dominicus – Cuello 1988: 28.
Tachybaptus dominicus – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported a femur, a tibiotarsus, and a coracoid from Lapa da Escrivânia V, a large tarsometatarsus from Lapa da Escrivânia IX, a large femur from Lapa da Escrivânia XI, and a humerus from an unknown locality. He noted that some might differ slightly from recent bones, probably individual variation, and two recent skeletons also diverge a little.
Podilymbus Lesson
304. Podilymbus podiceps (Linnaeus)
Quaternary – MG, RJ, RS
Podilymbus ( antarcticus vel podiceps , si sp. dist.) – Winge 1887: 25.
Podilymbus antarcticus – Goeldi 1894: 603.
Podilymbus antarcticus oder podiceps – Lambrecht 1933: 732.
Podilymbus podiceps – Brodkorb 1963: 230.
Podilymbus podicipes – Mones 1986: 76.
Podilymbus podiceps – Cuello 1988: 28.
Podilymbus podiceps – Nascimento and Silveira 2020: 490.
Winge ( 1887 ) reported the remains of at least three individuals from Lapa da Escrivânia V, represented by the posterior part of a mandibular branch, scapula, coracoid, humerus, synsacrum, femur, and tarsometatarsus. He associated them with Podilymbus antarcticus (= Podilymbus podiceps antarcticus ) or P. podiceps if they were de facto different taxa (treated as different subspecies by Pacheco et al. 2021 ). Only the tarsometatarsus was determined by comparison, with only bones from the skin of a male available from P. antarcticus . The tarsometatarsus is very similar to P. antarcticus , only slightly smaller, and differs from that of a P. podiceps in being slightly larger and narrower in the proximal end. The other bones match the tarsometatarsus when compared with the equivalents in Podiceps and Tachybaptus .
Podicidiformes [sic] – Kneip et al. 1989a: 126.
Podilymbus podiceps antarcticus – Kneip et al. 1989b: 662.
Podilymbus podiceps antarcticus [in part] – Kneip et al. 1994: 48.
Podicipedidae (“mergulhão”) [in part] – Kneip et al. 1995: 7.
Podicipedidae (“mergulhão”) [in part] – Kneip et al. 1997: 19, 37.
Podicipedidae (“mergulhão”) [in part] – Magalhães et al. 2001: 59.
Kneip et al. ( 1989b , 1994 ) reported three bones of at least one individual from layer I (3,800±190 years BP) and four bones of at least one individual from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Kneip et al. ( 1994 ) reported six bones of at least two individuals from layer III (1,810±40 years BP) of Sambaqui da Pontinha in Saquarema, Rio de Janeiro.
Podylimbus podiceps [sic] – Oliveira 2006: 318.
Oliveira ( 2006 ) reported three bones belonging to at least two individuals from the RS-RG-48 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Columbiformes Latham
Columbidae Leach
305. cf. Columbidae
Rosa ( 2004 ) reported a bone from the Paranaíba phase square 16H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
306. Columbidae indet. 1
Winge ( 1887 ) mentioned a small-sized columbid from Lapa do Capão Seco, among many fragile, hard-to-determine bones. He did not specify the skeletal elements found and thought it could represent some of the species already mentioned in his study.
307. Columbidae indet. 2
Quaternary – MG
Chamæpelia (et g. aff.) sp. 3 (fere certe): ut videtur: Ch. griseola vel Ch. ( Columbula ) campestris ; Ch. talpacoti ; Ch. ( Scardafella ) squamosa [in part] – Winge 1887: 39.
Chamaepelia campestris – Goeldi 1894: 386.
Chamoepelia sp. (“cfr. griseola , campestris , talpacoti und Scardafella squamosa ”) [in part] – Lambrecht 1933: 769.
Uropelia campestris (?) – Brodkorb 1971: 198.
Columbigallina passerina (?) – Brodkorb 1971: 198.
Uropelia campestris (?) – Brodkorb 1971: 198.
Columbigallina passerina [ Chamaepelia sp. [in part]] – Mones 1986: 93.
Uropelia campestris [ Chamaepelia sp. [in part]] – Mones 1986: 93–94.
Columbina passerina – Cuello 1988: 51.
Uropelia campestris – Cuello 1988: 52.
Columbidae indet. 2 – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported remains representing at least two species, probably three or even four, from Lapa da Escrivânia V except for a tibiotarsus from Lapa da Escrivânia XI associated with Columbina talpacoti . He divided them into four sets and noted that two Columbina talpacoti and two Columbina squammata skeletons were available for comparison. The first set comprises two small humeri, shorter and thinner than the other sets, which may represent Columbina passerina griseola or Uropelia campestris .
308. Columbidae indet. 3
Quaternary – MG
Chamæpelia (et g. aff.) sp. 3 (fere certe): ut videtur: Ch. griseola vel Ch. ( Columbula ) campestris ; Ch. talpacoti ; Ch. ( Scardafella ) squamosa [in part] – Winge 1887: 39.
Chamoepelia sp. (“cfr. griseola , campestris , talpacoti und Scardafella squamosa ”) [in part] – Lambrecht 1933: 769.
Chamaepelia sp. [in part] – Mones 1986: 93.
Columbidae indet. 3 – Nascimento and Silveira 2020: 492.
The second set of small columbid bones from Lapa da Escrivânia V defined by Winge ( 1887 ) is composed of elements larger than the first set, but still smaller than in Columbina talpacoti . It is represented by the humerus, ulna, radius, and tarsometatarsus (of which some may belong to the first set).
309. Columbidae indet. 4
Carvalho ( 1984 ) reported columbid remains among the material discovered in 1978 at the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
310. Columbidae indet. 5 (spp.?)
Carvalho et al. ( 1989 ) reported columbid remains from Furna do Estrago in Brejo da Madre de Deus, Pernambuco, collected by the staff of UNICAP. Lima ( 1992 ) reported the columbid remains from the site to consist of 469 bones.
311. Columbidae indet. 6
Late Holocene – SC
“pomba” – Schmitz and Verardi 1996: 143.
Schmitz and Verardi ( 1996 ) reported columbid remains from the Cabeçudas site in Itajaí, Santa Catarina.
312. Columbidae indet. 7
Rosa ( 2004 ) reported a bone from the Paranaíba phase square 18I of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
313. Columbidae indet. 8
Jacobus and Rosa ( 2013 ) reported columbid remains from the Sangão (RS-S-327) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul. Previously, this record was briefly mentioned by Dias and Jacobus ( 2003 ).
314. Columbidae indet. 9
Jacobus ( 2004 ) reported columbid remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
315. Columbidae indet. 10
Holocene – RS
Columbidae – Jacobus and Rosa 2013: 245.
Jacobus and Rosa ( 2013 ) reported columbid remains from the Deobaldino (RS-S-395) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
316. Columbidae indet. 11
Holocene – RS
Columbidae – Jacobus and Rosa 2013: 247.
Jacobus and Rosa ( 2013 ) reported columbid remains from the Schneider (RS-C-14) rock shelter in São Sebastião do Caí, Rio Grande do Sul.
317. Columbidae indet. 12
Holocene – RS
Columbidae – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported columbid remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
318. Columbidae indet. 13
Early Holocene – RS
Columbidae – Jacobus and Rosa 2013: 250.
Jacobus and Rosa ( 2013 ) reported columbid remains from the Garivaldino (RS-TQ-58) rock shelter in Montenegro, Rio Grande do Sul. The material is deposited in the CEPA collection.
Columbinae Leach
Patagioenas Reichenbach
319. cf. Patagioenas sp.
Late Holocene – RS
cf. Columba sp. – Rosa 2006b: 232.
Rosa ( 2006b ) reported a bone from level 4 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
320. Patagioenas picazuro (Temminck)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
321. Patagioenas cayennensis (Bonnaterre)
Quaternary – MG, RJ
Columba rufina – Winge 1887: 38.
Chloroenas rufina – Goeldi 1894: 386.
Columba rufina – Lambrecht 1933: 768.
Columba cayennensis – Brodkorb 1971: 197.
Columba cayennensis [ Columba rufina ] – Mones 1986: 93.
Columba cayennensis – Cuello 1988: 49.
Patagioenas cayennensis – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, somewhat larger than in a recent skeleton, represented by the scapula, coracoid, humerus, ulna, carpometacarpus, and femur. From Lapa da Escrivânia XI, there is the front part of a sternum, larger and slightly divergent, but still considered to belong to this species. There is also an ulna of recent age [28] .
Columba cayennensis – Bueno 2007: 34.
Bueno ( 2007 ) mentioned this species among avian remains from sambaquis in the APA de Massambaba, coastal Rio de Janeiro.
322. Patagioenas plumbea (Vieillot)
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, represented by the coracoid, humerus, ulna, femur, tibiotarsus, and tarsometatarsus. Additionally, there is a carpometacarpus and part of a skull from an unknown locality.
Geotrygon Gosse
323. Geotrygon montana (Linnaeus)
Quaternary – MG
Geotrygon montana – Winge 1887: 40.
Oropelia ( Geotrygon ) montana – Goeldi 1894: 386.
Geotrygon montana – Lambrecht 1933: 769.
Geotrygon montana – Brodkorb 1971: 199.
Geotrygon montana – Mones 1986: 93.
Geotrygon montana – Cuello 1988: 52.
Geotrygon montana – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported a tibiotarsus from Lapa da Escrivânia V, which he noted corresponds very closely to this species with only some individual variation and is quite distinct from Leptotila verreauxi . He also reported a coracoid and a femur of recent age [28] , which, when compared with a recent skeleton, are somewhat longer and slightly shorter, respectively.
Leptotila Swainson
324. Leptotila sp. 1
Late Holocene – RJ
“Provavelmente Leptotila rufaxila ou Leptotila verreauxi ” [sic] – Lima and Silva 1984: 30.
Lima and Silva ( 1984 ) reported remains possibly referable to Leptotila rufaxilla or Leptotila verreauxi from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. The material is represented by a left humerus, a right femur, and a left femur, all complete.
325. Leptotila spp. 2
Quaternary – MG
Leptotila spp. – Sales 2003: 214.
Sales ( 2003 ) reported four specimens among the bones and mummified carcasses from Zona II of Lapa do Rezar in Itacarambi, Minas Gerais.
326. Leptotila sp. 3
Holocene – PI
Leptotila sp. – Barbosa 2017: 110.
Barbosa ( 2017 ) reported this species from Toca do Alto da Serra do Capim in Guaribas, Piauí. These remains were not found in association with human burials or combustion structures.
327. Leptotila verreauxi Bonaparte
Quaternary – MG, PI, RJ
Engyptila ochroptera – Winge 1887: 39–40.
Leptoptila ochroptera ( chalcauchenia ) [sic] – Goeldi 1894: 386.
Engyptila ochroptera – Lambrecht 1933: 769.
Leptotila verreauxi – Brodkorb 1971: 199.
Leptotila verreauxi [ Engyptila ochroptera ] – Mones 1986: 93–94.
Leptotila verreauxi – Cuello 1988: 52.
Leptotila verreauxi – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported many bones from Lapa da Escrivânia V, with humeri notably numerous. They present a high degree of individual variation, particularly striking in some tarsometatarsi, but this was also observed in three recent skeletons. There are also a humerus from Lapa da Escrivânia XI, some bones from “various caves” [26] , and two bones of recent age [28] . The total material is represented by the scapula, coracoid, humerus, ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus.
Leptotila verreauxi – Bueno 2007: 34.
Bueno ( 2007 ) mentioned this species among avian remains from sambaquis in the APA de Massambaba, coastal Rio de Janeiro.
Leptotila verreauxi – Moraes 2014: 46, 60, 25.
Moraes ( 2014 ) reported this species from the uppermost layers of sector A of Toca da Janela da Barra do Antonião. The remains figure among material collected between 1988 and 1990 and are deposited in the Laboratório de Vestígios Orgânicos of FUMDHAM.
Only post-cranial remains were reported. About 90% of the analyzed humeri presented a dark coloration, which suggests they were burned. Some superficial fractures were also observed. A left humerus was depicted (labeled as 8257-7). A single ulna was analyzed, and it also had a dark coloration.
328. Leptotila rufaxilla (Richard & Bernard)
Late Holocene – RJ
Leptotila rufaxilla – Bueno 2007: 34.
Bueno ( 2007 ) mentioned this species among avian remains from sambaquis in the APA de Massambaba, coastal Rio de Janeiro.
Zenaida Bonaparte
329. Zenaida auriculata (Des Murs)?
Quaternary – MG
Columbid. indet. (s. under Zenaida maculata ) [in part] – Winge 1887: 13.
Zenaida maculata (?) – Winge 1887: 39.
Zenaïda maculata [sic] – Goeldi 1894: 385.
Zenaida maculata (?) (“kann auch Chamoepelia – Scardafella – squamosa sein”) – Lambrecht 1933: 769.
Zenaidura auriculata (?) – Brodkorb 1971: 198.
Zenaidura auriculata [ Zenaida maculata ?] – Mones 1986: 94.
Zenaida auriculata – Cuello 1988: 50.
Zenaida auriculata ? – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported four humeri from Lapa da Escrivânia V. They vary in size and are smaller than in Zenaida auriculata , but still very similar to it, and well-differentiated from Paraclaravis geoffroyi and Geotrygon montana . Additionally, there is a similar humerus from Lapa da Escrivânia XI, but it is even smaller, closer in size to that of Columbina squammata than that of Z. auriculata .
330. Zenaida auriculata (Des Murs)
Olson ( 1981 ) reported fossilized remains of Zenaida auriculata noronha found during his 1973 expedition on the Fernando de Noronha Archipelago, Pernambuco. They indicate the occurrence of this subspecies on the islands is not new. This insular taxon is the same found in parts of northeastern Brazil, probably due to the frequent inflows of individuals from the continent.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least eight adults and one young individual are present in the material.
Claravinae Richmond
Claravis Oberholser
331. Claravis pretiosa (Ferrari-Pérez)?
Quaternary – MG
Peristera cinerea ? – Winge 1887: 39.
[ Peristera ] cinerea – Goeldi 1894: 386.
Peristera cinerea (?) – Lambrecht 1933: 769.
Claravis pretiosa – Brodkorb 1971: 198.
Claravis pretiosa [ Peristera cinerea ?] – Mones 1986: 93–94.
Claravis pretiosa – Cuello 1988: 51.
Claravis pretiosa ? – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported some humeri from Lapa da Escrivânia V. They are considerably smaller than in Paraclaravis geoffroyi but morphologically equivalent, and he noted they almost certainly belong to Claravis pretiosa .
Paraclaravis Sangster, Sweet & Johnson
332. Paraclaravis geoffroyi (Temminck)
Quaternary – MG
“un Perroquet ( Psittacus )” [?; in part] – Gervais 1844a: 294.
Psittacus [in part] – Lund (in Winge 1887 ): 39.
Peristera geoffroyi – Winge 1887: 39.
Peristera Geoffroyi – Goeldi 1894: 385.
Peristera Geoffroyi – Lambrecht 1933: 769.
Psittacus sp. [?; in part] – Lambrecht 1933: 770.
Claravis godefrida – Brodkorb 1971: 199.
Claravis godefrida [ Peristera geoffroyi ] – Mones 1986: 93–94.
Claravis godefrida – Cuello 1988: 51.
Paraclaravis geoffroyi – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported some humeri from Lapa da Escrivânia V, which he noted are easily distinguishable from other genera, and a humerus from a “cave near Sumidouro” (other than Lapa da Lagoa do Sumidouro), determined as “ Psittacus ” in Lund’s catalog.
Columbina Spix
333. Columbina minuta (Linnaeus)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least five adults and one young individual are present in the material.
334. Columbina cf. talpacoti (Temminck)
Guérin et al. ( 1996 ) reported material comparable to this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
335. Columbina talpacoti (Temminck)
Quaternary – MG, RJ
Chamæpelia talpacoti [in part] – Winge 1887: 13.
Chamæpelia (et g. aff.) sp. 3 (fere certe): ut videtur: Ch. griseola vel Ch. ( Columbula ) campestris ; Ch. talpacoti ; Ch. ( Scardafella ) squamosa [in part] – Winge 1887: 39.
[ Chamaepelia ] Talpacoti – Goeldi 1894: 386.
Chamoepelia sp. (“cfr. griseola , campestris , talpacoti und Scardafella squamosa ”) [in part] – Lambrecht 1933: 769.
Columbigallina talpacoti – Brodkorb 1971: 198.
Columbigallina talpacoti [ Chamaepelia sp.” [in part]] – Mones 1986: 93.
Columbina talpacoti – Cuello 1988: 51.
Columbina talpacoti – Nascimento and Silveira 2020: 493.
The third set of small columbid bones established by Winge ( 1887 ) corresponds to Columbina talpacoti and is represented by two humeri and three ulnae from Lapa da Escrivânia V and a tibiotarsus from Lapa da Escrivânia XI.
Columbina talpacoti – Bueno 2007: 34.
Bueno ( 2007 ) mentioned this species among avian remains from sambaquis in the APA de Massambaba, coastal Rio de Janeiro.
336. Columbina squammata (Lesson)
Quaternary – MG
Chamæpelia (et g. aff.) sp. 3 (fere certe): ut videtur: Ch. griseola vel Ch. ( Columbula ) campestris ; Ch. talpacoti ; Ch. ( Scardafella ) squamosa [in part] – Winge 1887: 39.
Scardafella squamosa – Goeldi 1894: 386.
Chamoepelia sp. (“cfr. griseola , campestris , talpacoti und Scardafella squamosa ”) [in part] – Lambrecht 1933: 769.
Scardafella squammata – Brodkorb 1971: 198.
Scardafella squammata [ Chamaepelia sp.” [in part]] – Mones 1986: 93–94.
Scardafella squammata – Cuello 1988: 52.
Columbina squammata – Nascimento and Silveira 2020: 493.
The fourth set of small columbid bones established by Winge ( 1887 ) from remains from Lapa da Escrivânia V comprises four humeri, a carpometacarpus, and a tibiotarsus. They correspond to Columbina squammata , only slightly smaller, but somewhat larger than Columbina talpacoti .
337. Columbina picui (Temminck)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Cuculiformes Wagler
Cuculidae Leach
338. Cuculidae indet.
Rosa ( 2004 ) reported a bone from the Paranaíba phase square 14H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
Crotophaginae Swainson
Crotophaga Linnaeus
339. Crotophaga ani Linnaeus
Winge ( 1887 ) reported four humeri and a mandibular branch from Lapa da Escrivânia V.
Taperinae Verheyen
Tapera Thunberg
340. Tapera naevia (Linnaeus)
Quaternary – MG
Diplopterus nævius – Winge 1887: 45.
Diplopterus naevius – Goeldi 1894: 247.
Diplopterus naevius – Lambrecht 1933: 769.
Tapera naevia – Brodkorb 1971: 206.
Tapera naevia [ Diplopterus naevius ] – Mones 1986: 94.
Tapera naevia – Cuello 1988: 55.
Tapera naevia – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, represented by the coracoid, humerus, ulna, carpometacarpus, femur, and tarsometatarsus. Only bones from taxidermied specimens were available for comparison and the coracoid and femur were not determined by this method, as he noted they are quite distinct from the corresponding bones of other living cuculids found in the region.
Cuculinae Leach
Piaya Lesson
341. Piaya cayana (Linnaeus)
Quaternary – MG
Coccyzus cajanus [in part] – Lund 1841d: 18.
Coccyzus ? [in part] – Giebel 1846: 313.
Coccyzus [in part] – Giebel 1847: 19.
Coccyzus [in part] – Pictet 1853: 414.
“un Coua ( Coccyzus de Vieillot)” [in part] – Liais 1872: 303.
Coccyzus aff. cayano [in part] – Lund (in Winge 1887 ): 45.
Pyrrhococcyxcayanus – Winge 1887: 45.
Piaya cayana – Goeldi 1894: 247.
Coccyzus sp. (“angeblich aus den Knochenhöhlen Brasiliens”) [in part] – Lambrecht 1933: 769.
Pyrrhococcyx cayanus – Lambrecht 1933: 769.
Piaya cayana – Brodkorb 1971: 206.
Piaya cayana [ Pyrrhococcyx cayanus ] – Mones 1986: 94.
Piaya cayana – Cuello 1988: 54.
Piaya cayana – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported six humeri and a fragment of carpometacarpus from Lapa da Escrivânia V, and an incomplete humerus (“ Coccyzus aff. cayano ” in Lund’s catalog) from “a cave in Mocambo”.
Piaya cayana sp. – Sales 2003: 214.
Sales ( 2003 ) reported a specimen among the bones and mummified carcasses from Zona II of Lapa do Rezar in Itacarambi, Minas Gerais.
Nyctibiiformes Yuri, Kimball, Harshman, Bowie, Braun, Chojnowski, Hackett, Huddleston, Moore, Reddy, Sheldon, Steadman, Witt & Braun
Nyctibiidae Chenu & Des Murs
Nyctibius Vieillot
342. Nyctibius sp.
Quaternary – MG
Nyctibius sp. e mediis, fere certe N. jamaicensis – Winge 1887: 43–44.
“[provavel a identidade com] uma Nyctibius de tamanho mediano” – Goeldi 1894: 248.
Nyctibius sp. – Lambrecht 1933: 774.
“ Nyctibius Vieillot 1816 sp.” – Fisher 1967: 750.
Nyctibius griseus – Brodkorb 1971: 236.
Nyctibius griseus – Mones 1986: 96.
Nyctibius griseus – Cuello 1988: 57.
Nyctibius griseus – Sick 1984a: 334.
Nyctibius griseus – Sick 1993: 300.
Nyctibius griseus – Sick 1997: 408.
Nyctibius sp. – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported the proximal end of a left carpometacarpus from Lapa da Escrivânia V, and a right carpometacarpus lacking the upper part of recent age [28] . He noted that both fragments appear to belong to the same species, being much smaller and somewhat divergent from Nyctibius aethereus , and postulated they might belong, due to their measurements, to Nyctibius jamaicensis , since the other smaller species are too small.
Brodkorb ( 1971 ) listed the material as Nyctibius griseus , since N. jamaicensis does not occur in Brazil.
343. Nyctibius griseus (Gmelin)
Pleistocene – BA
Nyctibius griseus – Silva et al. 2012b: 368.
Silva et al. ( 2012b ) reported three carpometacarpi, two humeri, and a coracoid, representing six individuals, from Gruta dos Brejões in Morro do Chapéu, Bahia. It figures among material collected in 1980 and 1984 by the staff of the Laboratório de Paleontologia of MCL PUC Minas and was determined by Herculano Alvarenga.
Caprimulgiformes Ridgway
Caprimulgidae Vigors
344. Caprimulgidae indet. 1
Quaternary – MG
Caprimulgus [?] – Gervais 1844a: 294.
Caprimulgus [?] – Giebel 1846: 312.
Caprimulgus [?] – Giebel 1847: 8.
Caprimulgus [?] – Pictet 1853: 413.
“les Engoulevents” [?] – Liais 1872: 303.
Caprimulgus – Lund (in Winge 1887 ): 42.
G. sp. indet., forte Hydropsalis forcipata – Winge 1887: 42–43.
“provavel a identidade com Hydropsalis forcipata ” – Goeldi 1894: 248.
cfr. Hydropsalis forcipata – Lambrecht 1933: 773.
Cypselus ? affinis (“ Cypselus und Caprimulgus nach Lund in den Knochenhöhlen Brasiliens”) [?; in part] – Lambrecht 1933: 774.
Hydropsalis brasiliana [in part] – Brodkorb 1971: 235.
Hydropsalis [in part?] – Sick 1984a: 336.
Hydropsalis brasiliana [in part?] – Mones 1986: 96.
Hydropsalis [in part?] – Sick 1993: 303.
Hydropsalis [in part?] – Sick 1997: 412.
Caprimulgidae indet. 1 – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported a humerus (determined as “ Caprimulgus ” in Lund’s catalog) from a cave near Mocambo, quite similar to Hydropsalis torquata , but much larger, with matching dimensions to Hydropsalis forcipata . He noted that the similarity to Nyctidromus is significant, but no species with these dimensions are known. The humerus of Antrostomus rufus is much smaller and different in shape. Additionally, in the small details that differentiate it from H. torquata , it is somewhat similar to Podager , but much smaller than Podager nacunda .
345. Caprimulgidae indet. 2
Winge ( 1887 ) reported humeri in considerable number from Lapa da Escrivânia V. He noted they are generally smaller than in Hydropsalis torquata , from which a clear boundary could not be defined, but some fragments must belong to a different species, as some differences between the bones exist. Two of these humeri are quite peculiar in their morphology. There is less similarity to Antrostomus rufus than to H. torquata and Nyctidromus albicollis , but the bones may belong to a smaller species of Antrostomus . Winge used the contemporary definition of the genus and noted that, of these smaller species, Antrostomus ocellatus (= Nyctiphrynus ocellatus ) lives today in the region, and they could hardly belong to Chordeiles minor and Podager nacunda due to morphological differences.
Nyctidromus Gould
346. Nyctidromus albicollis (Gmelin)
Quaternary – MG
Nyctidromus albicollis – Winge 1887: 42.
Nyctidromus albicollis – Goeldi 1894: 248.
Nyctidromus albicollis – Lambrecht 1933: 773.
Nyctidromus albicollis – Brodkorb 1971: 235.
Nyctidromus – Sick 1984a: 336.
Nyctidromus albicollis – Mones 1986: 96.
Nyctidromus albicollis – Cuello 1988: 57.
Nyctidromus – Sick 1993: 303.
Nyctidromus – Sick 1997: 412.
Nyctidromus albicollis – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported several humeri from Lapa da Escrivânia V, well-differentiated from each other.
Hydropsalis Wagler
347. cf. Hydropsalis sp.
Guérin et al. ( 1996 ) reported material comparable to this genus from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
348. Hydropsalis anomala (Gould)
Quaternary – MG
Eleothreptus anomalus – Winge 1887: 42.
Eleothreptus anomalus – Goeldi 1894: 248.
Eleothreptus anomalus – Lambrecht 1933: 773.
Eleothreptus anomalus – Brodkorb 1971: 235.
Eleothreptus anomalus – Mones 1986: 96.
Eleothreptus anomalus – Cuello 1988: 58.
Hydropsalis anomala – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported two humeri from Lapa da Escrivânia V, one left and one right, and noted they are slightly smaller than those observed in a recent skeleton.
349. Hydropsalis torquata (Gmelin)
Quaternary – MG
Hydropsalis torqvata – Winge 1887: 43.
Hydropsalis torquata – Goeldi 1894: 248.
Hydropsalis torquata – Lambrecht 1933: 773.
Hydropsalis brasiliana [in part] – Brodkorb 1971: 235.
Hydropsalis [in part?] – Sick 1984a: 336.
Hydropsalis brasiliana [ Hydropsalis torqvata ] [in part?] – Mones 1986: 96.
Hydropsalis brasiliana – Cuello 1988: 57.
Hydropsalis [in part?] – Sick 1993: 303.
Hydropsalis [in part?] – Sick 1997: 412.
Hydropsalis torquata – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported several humeri from Lapa da Escrivânia V and some bones of recent age [28] represented by the mandible, sternum, scapula, ulna, and femur, which can be attributed to this species, although they show some differences. He noted that Nyctidromus albicollis is larger and divergent.
Apart from this material’s description and the remains he defined as G. sp. indet. ( Antrostomus , sp. e parvis, ?) (see Caprimulgidae indet. 2), Winge reported the existence of several bones from the same cave (Lapa da Escrivânia V) in addition to the humeri, represented by the coracoid, ulna, femur, and tibiotarsus. Part of these roughly match H. torquata , while part is too small and slightly divergent, without clear boundaries.
Apodiformes Peters
Apodidae Olphe-Galliard
Streptoprocne Oberholser
350. Streptoprocne zonaris (Shaw)
Quaternary – MG, PI
Cypsellus collaris [in part] – Lund 1841d: 18.
Cypselus ? [?; in part] – Giebel 1846: 313.
Cypselus [in part] – Giebel 1847: 8.
Cypselus collaris [in part] – Pictet 1853: 413.
“une autre [espèce] de celui des hirondelles” [?; in part] – Liais 1872: 303.
Cypsellus aff. collari [in part] – Lund (in Winge 1887 ): 44.
Chætura zonaris – Winge 1887: 44.
Chaetura zonaris – Goeldi 1894: 248.
Chaetura zonaris – Lambrecht 1933: 773.
Cypselus ? affinis (“ Cypselus und Caprimulgus nach Lund in den Knochenhöhlen Brasiliens”) [in part] – Lambrecht 1933: 774.
Streptoprocne zonaris – Brodkorb 1971: 240.
Streptoprocne zonaris – Sick 1984a: 345.
Streptoprocne zonaris [ Chaetura zonaris ] – Mones 1986: 97.
Streptoprocne zonaris – Cuello 1988: 58.
Streptoprocne zonaris – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported several individuals (including some very young) from Lapa da Escrivânia V, two bones from “various caves” [26] , and some bones from an unknown locality, including four humeri determined as “ Cypsellus aff. collari ” in Lund’s catalog. The total material is represented by the coracoid, sternum (anterior part), humerus (in large numbers), ulna, radius, carpometacarpus, the first phalanx of the major digit of the wing, femur, tibiotarsus, and tarsometatarsus.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Trochilidae Vigors
351. Trochilidae indet. 1
While living in Brazil, the French scientist Emmanuel Liais (1826–1900) found a well-preserved sternum among the bones of Lapa do Gambá in the Lagoa Santa region during his first voyage (Liais 1872 ). He believed it belonged to a hummingbird slightly larger than Trochilus minimus (= Mellisuga minima ) due to its shape and size. Ruschi ( 1979: 32), on the possible identity of the specimen, commented: “as in this locality some species of slightly larger size still live today, such as: Calliphlox a. amethystina Boddaert, 1831; Heliactin cornuta Wied, 1821; Chlorostilbon aureoventris pucherani Bourcier and Mulsant, 1848 and Amazilia versicolor kubitschecki Ruschi, 1959, it is difficult to determine the fragment without a comparative examination”. We contacted the staff of the Muséum Emmanuel Liais, in Cherbourg-en-Cotentin, France, about the possible whereabouts of this material, but it could not be located.
352. Trochilidae indet. 2
Quaternary – MG
G. sp. indet. e majoribus – Winge 1887: 44.
“uma especie, combinando no tamanho e na configuração com o Clytolaema rubinea ” [sic] – Goeldi 1894: 248.
sp. indet. größere Formen – Lambrecht 1933: 774.
“Undetermined species” – Storer 1960: 79.
Gen. and sp. indet. – Fisher 1967: 750.
Clytolaema rubricauda – Brodkorb 1971: 243.
Clytolaema rubricauda – Mones 1986: 97.
Clytolaema rubricauda – Cuello 1988: 58.
Clytolaema rubricauda – Unwin 1993: 729.
Trochilidae indet. 2 – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a carpometacarpus from Lapa da Escrivânia V of a large species, somewhat larger than that observed in Heliodoxa rubricauda .
Brodkorb ( 1971 ), most likely following Winge’s comment, listed the material under this name, associating it with the locality “Lapa da Escrivânia?”.
Phaethornithinae Jardine
Phaethornis Swainson
353. Phaethornis pretrei (Lesson & Delattre)?
Winge ( 1887 ) reported a carpometacarpus and a fragment of the anterior part of a sternum, both of recent age, without attributing them to a specific locality. They belong to a smaller form than the indeterminate trochilid [30] from Lapa da Escrivânia V (see above). The size and shape correspond to Phaethornis pretrei , but whether it belongs to this species could not be defined.
354. Phaethornis pretrei (Lesson & Delattre)
Quaternary – MG
Phaethornis pretrei sp. – Sales 2003: 214.
Sales ( 2003 ) reported a specimen among the bones and mummified carcasses from Zona II of Lapa do Rezar in Itacarambi, Minas Gerais.
Trochilinae Vigors
Chlorestes Reichenbach
355. Chlorestes notata (Reich)
Quaternary – MG
Chlorestes notatus sp. – Sales 2003: 214.
Sales ( 2003 ) reported a specimen among the bones and mummified carcasses from Zona II of Lapa do Rezar in Itacarambi, Minas Gerais.
Opisthocomiformes Sclater
Opisthocomidae Swainson
†Hoazinavis Alvarenga, Mayr & Mourer-Chauviré
356. † Hoazinavis lacustris Alvarenga, Mayr & Mourer-Chauviré
Late Oligocene/Early Miocene – SP
Fig. 32 A
Type locality: Tremembé Formation, Taubaté Basin, State of São Paulo, Brazil.
Etymology: Hoazinavis , a combination of the vernacular name hoatzin and the Latin avis , and the Latin lacustris , referring to the lake deposits of the Tremembé Formation.
Hoazinavis lacustris gen. et sp. nov. [new genus and species] – Mayr et al. 2011b: 961–966, 1D, E, H, I, M, N.
Hoazinavis lacustris – Mayr 2022: 93, 6.1B.
Hoazinavis lacustris – Carmo et al. 2024: 5.
Mayr et al. ( 2011b ) described this new genus and species from material collected in 2008 by Herculano Alvarenga (Pivetta 2011 ) in the Tremembé Formation. It was considered then the unmistakably oldest and smallest opisthocomiform.
The material (MHNT-VT-5332, holotype) consists of a complete right humerus, the omal end of a right coracoid, and the cranial end of a right scapula, all from the same adult individual.
In the same study, Mayr et al. reclassified Namibiavis senutae , a species from the early Miocene of Namibia (Mourer-Chauviré 2003 ), from Idiornithidae to Opisthocomiformes, and through phylogenetic analysis placed it as the sister of a clade formed by Hoazinavis and Opisthocomus .
The remains of both species indicate they were weak distance flyers, just like the living Opisthocomus hoazin , which, due to the proportionally large crop, developed a modified sternal keel and pectoral musculature to accommodate it. Whether and to what degree the two fossil species were already folivorous cannot be stated, but a large crop considerably affects the morphology of the pectoral girdle bones. Due to the similarities of the known elements with those of O. hoazin , Mayr et al. believed it is likely that at least H. lacustris had already evolved a large crop and folivory at some level, whose development in the only living species suggests a very long evolutionary history of specialized feeding.
The discovery of Namibiavis in Africa shows that the current distribution of hoatzins is relictual and implies the distribution of stem Opisthocomiformes was the result of dispersion and not vicariance, given the fact that South America and Africa separated during the Cretaceous before the appearance of crown Neornithes in the fossil record. Because of this, Mayr et al. believed that, even if the Miocene hoatzins had better flight capabilities than the living ones and the existence of islands in the South Atlantic during the beginning of the Cenozoic is considered, the dispersion by the ocean (minimum distance of 1,000 km in a straight line during that time) through floating vegetation rafts is the most plausible explanation. The folivorous diet would favor these birds in these rafts, which come off the mouth of large rivers and can reach considerable sizes. Therefore, an African origin is more likely, as was the case with caviomorph rodents, platyrrhinous primates, and some South American amphibians and geckos, favored by paleocurrents and paleowinds. However, additional fossils and a better understanding of the phylogenetic relationships of the Opisthocomiformes are needed for the establishment of a hypothesis of dispersion direction. Additionally, Mayr ( 2014 ) reported a middle Miocene tarsometatarsus from Kenya (Maboko Formation) assigned to ? Namibiavis sp., which has an essentially modern morphology and indicates arboreal habits consistent with a folivorous diet.
The theory of a hoatzin origin in the Old World and their current relictual distribution were corroborated by the description of European fossils. Mayr and De Pietri ( 2014 ) described Protoazin parisiensis from the late Eocene of France, the oldest known representative of the group and the first found in the Northern Hemisphere. The material shows greater affinity with Opisthocomus and Hoazinavis than with Namibiavis . However, Mayr and De Pietri noted that the theory of an alternative dispersion route through the Northern Hemisphere lacks support with the absence of known fossils in North America.
From South America, the only other known fossil opisthocomiform is Hoazinoides magdalenae (Miller 1953 ), from the middle Miocene of Colombia (Villavieja Formation). It is slightly larger than O. hoazin , and, along with H. lacustris , indicate South American records outside the group’s current geographical distribution (Mayr et al. 2011b ).
Gruiformes Bonaparte
Aramidae Bonaparte
Aramus Vieillot
357. Aramus guarauna (Linnaeus)
Late Holocene – RS
Aramidae – Milheira et al. 2019: 45.
Milheira et al. ( 2019 ) reported aramid remains among the material from the sites PSG-02 and PSG-07 of the Pontal da Barra cerrito complex in Pelotas, Rio Grande do Sul. It likely represents Aramus guarauna as it is the only known recent species.
Rallidae Rafinesque
358. cf. Rallidae
Rosa ( 2009 ) reported 12 possible rallid bones (seven from square A6, two from C5, two from D6, and one from D7) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Remains from square A6 were detected in the site’s three occupation periods. The material is deposited in the CEPA collection.
359. Rallidae indet. 1
Quaternary – MG
G. sp. indet. (similis præcedenti [ Rallus nigricans ]) – Winge 1887: 26.
“ Rallus spec., parecido com a especie precedente [ Rallus ( A ) . nigricans ]” – Goeldi 1894: 559.
Rallus nigricans (?) [in part] – Lambrecht 1933: 760.
Ortygonax nigricans (?) [in part] – Brodkorb 1967: 133.
Rallus nigricans [in part] – Cuello 1988: 42.
Rallidae indet. 1 – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported as indeterminate a humerus from Lapa do Capão Seco, similar to the largest specimens of the species he presented just before this record in his list ( Pardirallus nigricans ). It presents a very large internal condyle.
360. Rallidae indet. 2
Quaternary – MG
Crex minuta [in part] – Lund 1841d: 18.
Crex ? [in part] – Giebel 1846: 311.
“einer der dort noch lebenden Cr. minuta auffallend ähnlichen Art” [in part] – Giebel 1847: 30.
Rallus [in part] – Pictet 1853: 420.
“les Rallus” [in part] – Liais 1872: 303.
Rallus minutus [in part] – Lund (in Winge 1887 ): 26.
Porzana sp. e minimis, non P. flaviventris – Winge 1887: 26–27.
“[ Porzana ] spec. (especie pequenissima)” – Goeldi 1894: 559.
Porzana sp. – Lambrecht 1933: 761.
“other species” – Brodkorb 1967: 134.
Porzana flaviventris – Mones 1986: 86.
Porzana flaviventer – Cuello 1988: 42.
Rallidae indet. 2 – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V (perhaps the most abundant bird in that site), a bone from Lapa da Escrivânia III, several bones from Lapa da Lagoa do Sumidouro, some bones from an unknown locality, and material of recent age [28] in large volume (including a humerus that Lund determined as “ Rallus minutus ”). These numerous materials indicate it was apparently one of the most common birds in the region, and the fact that neither Lund nor Reinhardt found it alive may be a consequence of its secretive habits. The total material is represented by several head parts, scapula, coracoid, furcula, sternum, humerus, ulna, radius, carpometacarpus, pelvis, femur, tibiotarsus, and tarsometatarsus. Winge noted that the bones do not belong to Porzana (= Laterallus ) flaviventer , which he defined as the only small species of the genus known in the region. The tarsometatarsus is slightly smaller than that of L. flaviventer , but the forearm bones are much larger, and, in their regard, Winge remarked it is necessary to consider the great individual variation observed in the tarsometatarsi and ulnae from the caves. One unusually short tarsometatarsus is similar to that of Porphyriops , but it was not possible to determine whether it is an individual variation or another species.
Mones ( 1986 ) listed the record as Porzana flaviventris and noted he followed Brodkorb ( 1967 ), who, at any rate, said these materials belong to another species.
361. Rallidae indet. 3
Winge ( 1887 ) reported several humeri from Lapa da Escrivânia V. They are relatively short and robust (he provided the length of five of them), and the size differences could indicate two species. The longer ones have the same length and general shape as those of Porphyriops melanops but are more robust, with wider ends. They are quite different from those of Porphyrio flavirostris . There is also a tarsometatarsus very similar to that of P. melanops but smaller, and different from all the others he could compare.
362. Rallidae indet. 4
Winge ( 1887 ) reported a tarsometatarsus of a small rallid from Lapa da Lagoa do Sumidouro, which hardly belongs to the other rallids he mentioned in his study.
363. Rallidae indet. 5
Carvalho ( 1984 ) reported rallid remains among the material discovered in 1978 at the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
364. Rallidae indet. 6
Guérin et al. ( 1996 ) reported material comparable to the genus “ Gallinule ” (= Gallinula ?) from Toca da Janela da Barra do Antonião.
365. Rallidae indet. 7
Late Holocene – SC
“saracura” – Schmitz and Verardi 1996: 143.
Schmitz and Verardi ( 1996 ) reported rallid remains from the Cabeçudas site in Itajaí, Santa Catarina.
366. Rallidae indet. 8
Jacobus and Rosa ( 2013 ) reported rallid remains from the Sangão (RS-S-327) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul. Previously, this record was briefly mentioned by Dias ( 2003 , 2004a ).
367. Rallidae indet. 9
Jacobus ( 2004 ) reported rallid remains from Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
368. Rallidae indet. 10 (spp.?)
Rosa ( 2004 ) reported indeterminate rallids from the GO-JA-01 rock shelter in Serranópolis, Goiás. From the Paranaíba phase (early Holocene), a bone from square 14H. From the Serranópolis phase (early–late Holocene), a bone from cut 1/2. Finally, from the Jataí phase (late Holocene), a bone from square 18H. The material is deposited in the IAP/Unisinos collection.
369. Rallidae indet. 11
Late Holocene – RS
Rallidae – Rosa 2006b: 231.
Rosa ( 2006b ) reported a bone from level 4 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
370. Rallidae indet. 12
Late Holocene – RS
Rallidae – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported rallid remains from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. They highlighted that these birds are exceptionally represented by some dozens of individuals. The material is deposited in the IAP/Unisinos collection.
371. Rallidae indet. 13
Jacobus and Rosa ( 2013 ) reported rallid remains from the Deobaldino (RS-S-395) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
372. Rallidae indet. 14
Holocene – RS
Rallidae – Jacobus and Rosa 2013: 247.
Jacobus and Rosa ( 2013 ) reported rallid remains from the Schneider (RS-C-14) rock shelter in São Sebastião do Caí, Rio Grande do Sul.
373. Rallidae indet. 15
Holocene – RS
Rallidae – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported rallid remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
374. Rallidae indet. 16
Quaternary – MG
Rallidae – Seersholm et al. 2021: 2067.
Seersholm et al. ( 2021 ) reported an indeterminate rallid among the bones collected by Lund in Lapa da Escrivânia V that were analyzed through bulk bone metabarcoding. It was also detected morphologically in one of the studied samples.
Rallus Linnaeus
375. † Rallus sp.
Holocene – PE
Fig. 34 D
“a medium-sized species of rail” – Olson 1977: 357.
Rallidae – Olson 1981: 483.
“saracura fóssil” – Sick 1984a: 250.
Rallidae – Alvarenga 1992: 254.
Rallidae – Alvarenga 1993a: 63–64.
“sub-fossil rail” – Sick 1993: 211.
Rallidae – Alvarenga 1997: 123.
“saracura fóssil” – Sick 1997: 296.
Rallus sp. – Carleton and Olson 1999: 55.
? Atlantisia sp. – Hume 2017: 116–117.
While theorizing on the absence of species of Atlantisia or another genus in South Atlantic islands, Olson ( 1973 ) speculated on the existence of a rallid species exterminated by man-introduced mammals in the Fernando de Noronha Archipelago ( Fig. 1 .19) or in the Trindade Island before specimens could be obtained. He noted that a search for subfossil remains on these territories could provide interesting results. During an expedition to the Fernando de Noronha Island in 1973, Olson ( 1977 , 1981 ) collected the remains of a medium-sized rallid in old beach dunes at the base of the Santo Antônio peninsula, along with other vertebrates, gastropods, and land crabs, of a probably late holocenic age. Most of the dune material was scattered and many specimens were well mineralized (Olson 1981 , Carleton and Olson 1999 ).
Most elements of the skeleton were found, associated with several individuals. It had reduced wings and may have been flightless but to a lesser extent than other species. Olson ( 1977 , 1981 ) initially postulated that it did not seem to be referable to any genera of mainland Brazil and might have been a form of Atlantisia , a genus now represented only by Atlantisia rogersi from Inaccessible Island in the Tristan da Cunha Archipelago (Taylor 1996 ). The remains were later associated with the genus Rallus (Carleton and Olson 1999 ).
The first mention of the archipelago’s fauna is found in the account of the disputed fourth voyage of the explorer Amerigo Vespucci (1454?–1512)—“ Lettera di Amerigo Vespucci delle Isole Nuovamente in Quattro Suoi Viaggi ” (1504), written in 1503, the year in which the discovery of the islands is generally attributed to (Olson 1981 , Carleton and Olson 1999 ). Vespucci described marine and land birds occurring in large numbers, “so guileless that they let themselves be taken in hand” and had “loaded a boat full of them”; he also mentioned reptiles and “very large rats” (Carleton and Olson 1999: 47). The latter likely represents the now-extinct Noronhomys vespucci , described by Carleton and Olson from material discovered by Olson on his expedition. After Vespucci’s account, no biological survey on the islands would be made until the arrival of naturalists J.C. Branner in 1876 and H.N. Ridley in 1887 (Carleton and Olson 1999 ). If the rail was among the terrestrial birds observed by Vespucci, the action of man in direct hunting, habitat alteration, and/or in the introduction of predatory species of both eggs and adults ( Noronhomys vespucci was interpreted as predominantly herbivore) had a devastating effect on this bird, which evolved in isolation, leading to its extinction shortly afterward, as happened with several other rallids on islands across the planet.
The complete description of the material, deposited at the Smithsonian Institution (Washington, D.C.) (Hume 2017 ), is still pending. Olson passed away in January 2021, but in 2015–2016 personal communications with us expressed that the description was something he should have attended to long ago. After revising his ideas on Atlantisia , he reasserted that the rail of Fernando de Noronha is most likely a derivate of Rallus , as are all the fossil rails from the North Atlantic found so far (Bermuda and the Macaronesian Islands of Azores and Madeira), so this widely dispersing genus probably made it across the Equator to Fernando de Noronha. The fossils were never depicted, but Olson commented that its bill is of medium length, not extremely elongated as in some species of Rallus , such as the larger ones, probably similar to the extinct Macaronesian Islands rallids eventually described by Alcover et al. ( 2015 ) but perhaps a little stouter. He also noted that almost none of the known fossil material from Fernando de Noronha has been cataloged or numbered yet. Additionally, regarding the age of the total material from the site, he noted that most of the fossils are indeed probably holocenic, although some, from their geological context, are almost certainly pleistocenic, but they are few and not particularly important.
On the absence of rallid remains in Trindade, Olson ( 1981 ) noted it is very unlikely that no endemic species existed there in the past, given the ample habitat and that these birds have successfully colonized all the other South Atlantic islands. Possibly, he did not find any remains because he was unlucky in his prospections, with the action of land crabs also reducing the chances of any carcass surviving long enough to be preserved, despite land crabs also occurring in Fernando de Noronha. The absence of resident Procellariiformes even in the fossil record of Fernando de Noronha may also be attributed to this predatory action on their nests, making reproduction very difficult or impossible.
Porphyrio Brisson
376. cf. Porphyrio martinica (Linnaeus)
Guérin et al. ( 1996 ) reported material comparable to the genus Porphyrula (= Porphyrio martinica ) from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
377. Porphyrio martinica (Linnaeus)?
Quaternary – MG
Porphyrio martinicus ? – Winge 1887: 27.
[ Porphyrio ] martinica – Goeldi 1894: 559.
Porphyrio sp. (“(?) martinicus ”) – Lambrecht 1933: 761.
Porphyrula martinica (?) – Brodkorb 1967: 135.
Porphyrula martinica [ Porphyrio martinicus ?] – Mones 1986: 86.
Porphyrula martinica – Cuello 1988: 43.
Porphyrio martinicus ? – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) associated, inconclusively, two humeri from Lapa da Escrivânia V with this species.
Laterallus Gray
378. Laterallus melanophaius (Vieillot)?
Quaternary – MG
Porzana melanophæa ? – Winge 1887: 26.
[ Porzana ] melanophaea – Goeldi 1894: 559.
Porzana sp. (“(?) melanocephala ”) – Lambrecht 1933: 761.
Porzana melanophaius – Brodkorb 1967: 134.
Porzana melanophaius ? – Mones 1986: 86.
Laterallus melanophaius – Cuello 1988: 43.
Laterallus melanophaius ? – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported two tibiotarsi and two tarsometatarsi from Lapa da Escrivânia V, and one tarsometatarsus of recent age [28] , similar to the remains he described as “ Porzana sp. and minimis, non P. flaviventris ” (see Rallidae indet. 2) but larger and slenderer. Of the species that live in the region, he believed they could only belong to Laterallus melanophaius , of which the tarsometatarsus of a taxidermied specimen corresponds in size. Additionally, several humeri from Lapa da Escrivânia V, similar in size to Porphyrio flavirostris but different in shape, probably belong to this species. Winge noted they are perhaps quite large regarding the other bones, but this does not prevent them from belonging to the same species.
Mustelirallus Bonaparte
379. Mustelirallus albicollis (Vieillot)
Quaternary – MG
Porzana albicollis – Winge 1887: 26.
Porzana ( Ortygometra ) albicollis – Goeldi 1894: 559.
Porzana albicollis – Lambrecht 1933: 761.
Porzana albicollis – Brodkorb 1967: 134.
Porzana albicollis – Mones 1986: 86.
Porzana albicollis – Cuello 1988: 42.
Mustelirallus albicollis – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) associated with this species bones from several individuals from Lapa da Escrivânia V. He noted they are very similar to Pardirallus nigricans , but some bones may belong to either a small representative of that species or to a large Mustelirallus albicollis .
Pardirallus Bonaparte
380. Pardirallus nigricans (Vieillot)
Quaternary – MG
Rallus nigricans (et. aff.) – Winge 1887: 26.
Rallus ( A ) . nigricans – Goeldi 1894: 559.
Rallus nigricans (?) [in part] – Lambrecht 1933: 760.
Ortygonax nigricans [in part] – Brodkorb 1967: 133.
Ortygonax nigricans [ Rallus nigricans ] – Mones 1986: 86.
Rallus nigricans [in part] – Cuello 1988: 42.
Pardirallus nigricans – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) associated with this species the bones of several individuals from Lapa da Escrivânia V but noted that some larger ones may belong to another taxon. Additionally, there is a bone of recent age [28] .
Aramides Pucheran
381. cf. Aramides sp.
Schmitz and Ferrasso ( 2011 ) reported the distal end of a tibiotarsus comparable to this genus from the Guarani site Itapiranga I (SC-U-1) in Itapiranga, Santa Catarina. The occurrence of this genus in the site was previously mentioned by Ferrasso and Schmitz ( 2010 ). The material was discovered in January 1957 and is deposited in the IAP/Unisinos collection.
382. Aramides sp. 1
Quaternary – MG
Aramides sp. ( cayennensis v. serracura ) – Winge 1887: 25–26.
Aramides spec. ( cayennensis ) – Goeldi 1894: 559.
Aramides cayanea (“oder A. serracura ”) – Lambrecht 1933: 760.
Aramides cajanea – Brodkorb 1967: 133.
Aramides cajanea – Mones 1986: 86.
Aramides cajanea – Cuello 1988: 42.
Aramides sp. – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported a humerus from “various caves”* [26] and a femur from an unknown locality as classifiable under Aramides cajaneus . However, he noted that, due to their significant similarity to Aramides saracura* and the lack of a significant number of specimens for comparison, it was not possible to determinate it at the species level.
Brodkorb ( 1967 ) and, following him, Cuello ( 1988 ) listed the material as from “Lapa da Escrivania?”, probably due to Winge ( 1887: 12) having listed the taxon under “I Lapa da Escrivania Nr. 5 mangle blot følgende” (“In Lapa da Escrivânia V only the following are missing”).
383. Aramides sp. 2
Dias ( 2004b ) and Rosa ( 2004 ) reported this genus from the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
384. Aramides sp. 3
Late Holocene – RS
Aramides sp. – Rosa 2006b: 231.
Rosa ( 2006b ) reported two bones from level 4 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
385. Aramides sp. 4
Middle Holocene – SC
Aramides sp. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) associated with this genus four bones of at least two individuals from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
386. Aramides cajaneus Statius Müller
Late Holocene – RJ
Aramides cajanea – Lima and Silva 1984: 29–30.
Lima and Silva ( 1984 ) reported remains from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. The material includes at least 13 individuals and is represented by a complete synsacrum; the humerus: three complete right ones, one almost complete, five almost complete left ones, part of a right diaphysis, and part of a left diaphysis; the ulna: two complete left ones and a distal left half; the femur: three left ones (two complete and one almost complete), four almost complete right ones, a partial diaphysis, the distal left half, and the proximal right half; the tibiotarsus: two complete right ones, a complete left one, two proximal right halves, two proximal right ends, a proximal left end, two partial diaphyses (one right), two proximal partial diaphyses (one right), a distal half of the left diaphysis, two right distal halves, three right distal parts, a left distal part, and a right distal fragment; and the tarsometatarsus: five left ones (three complete and two almost complete), a complete right one, a distal right half, and the proximal part of the right dyaphisis.
387. Aramides saracura (Spix)
Kneip et al. ( 1986 ) reported this species from layer 1 of Sambaqui Zé Espinho “A” in Rio de Janeiro, Rio de Janeiro, dated 1,510±160 years BP. The material, excavated in 1983, comprises a complete skull (Vogel 1987 ) and is deposited in the MN.
Porzana Vieillot
388. Porzana carolina (Linnaeus)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material. This species, which breeds in North America and winters in Central and northern South America, was previously not known to reach Brazil in the present until the first authentic record was published by Camacho and Accorsi ( 2016 ) for Rio de Janeiro.
Gallinula Brisson
389. Gallinula galeata (Lichtenstein)
Quaternary – MG
Gallinula galeata – Winge 1887: 27.
Gallinula galeata – Goeldi 1894: 559.
Gallinula galeata – Lambrecht 1933: 761.
Gallinula chloropus – Brodkorb 1967: 135.
Gallinula chloropus [ Gallinula galeata ] – Mones 1986: 86.
Gallinula chloropus – Cuello 1988: 43.
Gallinula galeata – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported a humerus and a tarsometatarsus from Lapa da Escrivânia V.
Charadriiformes Huxley
Charadriidae Leach
390. Charadriidae indet. 1
Late Holocene – RS
Charadriidae – Ferrasso et al. 2013: 225.
Ferrasso et al. ( 2013 ) reported an indeterminate charadriid humerus about 1.5 cm long from Sambaqui Arroio Seco V (RS-LN-285) in Arroio do Sal, Rio Grande do Sul.
391. Charadriidae indet. 2
Middle Holocene – SC
Charadriidae indet. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) reported two indeterminate charadriid bones of at least one individual from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
Vanellus Brisson
392. Vanellus sp.
Quaternary – MG
Vanellus aff. cayennensi , forma magna; Vanellus cayennensis et aff. [“Knogler af en Art eller Form, der staar ganske nær ved V. cayennensis og occidentalis , men er større”] – Winge 1887: 13, 27–28.
*“Vanellus cayennensis* (2 formas, uma grande e uma pequena)” [in part] – Goeldi 1894: 559.
Vanellus cayennensis [in part] – Lambrecht 1933: 763.
Belonopterus chilensis – Brodkorb 1967: 199.
Belonopterus chilensis – Mones 1986: 92.
Vanellus chilensis – Cuello 1988: 45.
Belonopterus chilensis – Cenizo and Agnolín 2007: 41.
Vanellus sp. – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported an almost complete right humerus (in two parts) from Lapa da Escrivânia V, which corresponds well to that of Vanellus chilensis (“ V. cayennensis or occidentalis ”) but is considerably larger and presents a longer deltoid crest. Additionally, there is the proximal end of another right humerus from Lapa da Escrivânia XI, similar to the previous one.
393. Vanellus chilensis (Molina)
Quaternary – GO, MG, MS
Vanellus cayennensis et aff. [“Knogler af Størrelse som hos V. cayennensis ”] – Winge 1887: 28.
*“Vanellus cayennensis* (2 formas, uma grande e uma pequena)” [in part] – Goeldi 1894: 559.
Vanellus cayennensis [in part] – Lambrecht 1933: 763.
Vanellus chilensis – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported the distal end of a tibiotarsus, a coracoid, and a fragment of the frontal and medial parts of the sternum of a young individual from Lapa da Escrivânia V. Despite small variations, these bones correspond well in size to Vanellus chilensis .
Rosa ( 2004 ) reported a bone from square 14H and another from the Paranaíba phase (early Holocene) square 18I of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
Vanellus chilensis – Schmitz et al. 2009: 342.
Schmitz et al. ( 2009 ) reported two bones from level 5 of the MS-MA-16A site in Corumbá, Mato Grosso do Sul.
Scolopacidae Rafinesque
394. Scolopacidae indet.
Winge ( 1887 ) reported the proximal part of an ulna of a bird similar to Tringa or the then valid Totanus from Lapa da Escrivânia V. It is insignificantly smaller than and slightly different from that of a Tringa melanoleuca , but this may be due to individual variation.
Arenariinae Stejneger
Calidris Merrem
395. Calidris cf. melanotos (Vieillot)
Quaternary – MG
Tringa maculata (fere certe) – Winge 1887: 28.
Tringa maculata – Goeldi 1894: 559.
Tringa maculata – Lambrecht 1933: 763.
Erolia melanotos – Brodkorb 1967: 196.
Erolia melanotos [ Tringa maculata ?] – Mones 1986: 91.
Calidris melanotos – Cuello 1988: 48.
Calidris cf. melanotos – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) associated, with uncertainty, a humerus from Lapa da Escrivânia V with this species. He noted that only a recent humerus lacking the proximal part was available for comparison but, even so, the similarity between the two bones was so pronounced they almost certainly belong to the same species.
396. Calidris cf. pusilla (Linnaeus)
Quaternary – MG
Ereunetes pusillus (fere certe) – Winge 1887: 28.
Ereunetes pusillus – Goeldi 1894: 559.
Ereunetes pusillus – Lambrecht 1933: 764.
Ereunetes pusillus – Brodkorb 1967: 196.
Ereunetes pusillus ? – Mones 1986: 91.
Calidris pusilla – Cuello 1988: 48.
Calidris cf. pusilla – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) associated, with uncertainty, a humerus from Lapa da Escrivânia V with this species. He noted that what he said about the material associated with Calidris melanotos also applies to this one, with the recent bone available for comparison only slightly more incomplete. The humerus is larger than in Calidris minuta and even larger than in Calidris temminckii , but much smaller than in Calidris bairdii and quite different from Actitis macularius .
Vanellus sp., Lapa da Escrivânia V: right humerus, caudal (A) and cranial (B) views; Lapa da Escrivânia XI: proximal end of the right humerus, caudal (C) and cranial (D) views (K.L. Hansen). Scale bar: 10 mm.
Scolopacinae Rafinesque
Gallinago Brisson
397. Gallinago cf. paraguaiae (Vieillot)
Quaternary – MG
Gallinago frenata (v. sp. aff.) – Winge 1887: 28.
Scolopax frenata – Goeldi 1894: 559.
Gallinago frenata (?) – Lambrecht 1933: 764.
Capella paraguaiae – Brodkorb 1967: 197.
Capella paraguaiae [ Gallinago frenata ] – Mones 1986: 91.
Gallinago gallinago paraguaiae – Cuello 1988: 47.
Gallinago cf. paraguaiae – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, represented by the frontal part of the sternum, humerus, ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus. Most bones are slightly larger and somewhat divergent than in a specimen of Gallinago paraguaiae , but, in general, they belong to a very similar form, which, likewise, had shorter and stronger legs and weaker wings than Gallinago gallinago . There are also marked differences between the fossil bones, particularly in the humeri. Furthermore, there is a coracoid from Lapa da Lagoa do Sumidouro.
Tringinae Rafinesque
Tringa Linnaeus
398. Tringa solitaria Wilson
Quaternary – MG
Totanus solitarius – Winge 1887: 28.
Totanus solitarius – Goeldi 1894: 559.
Totanus solitarius – Lambrecht 1933: 764.
Tringa solitaria – Brodkorb 1967: 195.
Tringa solitaria [ Totanus solitarius ] – Mones 1986: 91.
Tringa solitaria – Cuello 1988: 46.
Tringa solitaria – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) associated with this species a humerus and possibly an ulna from Lapa da Escrivânia V.
Jacanidae Chenu & Des Murs
Jacana Brisson
399. Jacana jacana (Linnaeus)
Quaternary – MG
Parra jacana – Winge 1887: 29.
Parra jaçanã [sic] – Goeldi 1894: 559.
Jacana jacana – Lambrecht 1933: 762.
Jacana spinosa – Storer 1960: 74.
Jacana spinosa – Brodkorb 1967: 202.
Jacana spinosa – Fisher 1967: 746.
Jacana spinosa – Olson 1976: 259.
Jacana spinosa [ Parra jacana ] – Mones 1986: 91.
Jacana jacana – Cuello 1988: 44.
Jacana jacana – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported three humeri and a carpometacarpus from Lapa da Escrivânia V and a carpometacarpus from Lapa da Lagoa do Sumidouro.
Laridae Rafinesque
400. Laridae indet. 1
Quaternary – MG
G. sp. indet. Larus (s. lat.) e minoribus, vel Sterna (s. lat.) e majoribus – Winge 1887: 29.
“um exemplar, de genero e especie desconhecidos, ou uma especie pequena de Larus , ou grande de Sterna ” – Goeldi 1894: 603.
Larus sp. (“kann auch Sterna sein”) – Lambrecht 1933: 766.
Pseudosterna degener (?) – Mercerat 1897: 238.
Laridae ( sensu lato ) indet. – Nascimento and Silveira 2020: 492.
Winge ( 1887 ) reported a carpometacarpus lacking the inferior part from Lapa da Escrivânia V, which may have belonged to a large tern or a small gull. It is similar to that of Rynchops niger but slightly smaller.
Mercerat ( 1897 ) raised the possibility that this material belongs to Pseudosterna degener , genus and species he erected for material of the Pleistocene of Argentina (Brodkorb 1967 ). However, his assumption was based on Winge’s texts only. In addition to being represented by different skeletal elements than those known from the larid of the Lagoa Santa region, the type material of both Pseudosterna degener and Pseudosterna pampeana , a second species described on the same occasion, were never illustrated and were subsequently lost (Olson 1985b , Mones 1986 ).
401. Laridae indet. 2
Late Holocene – RJ
Sterna (?) – Lima and Silva 1984: 30.
Lima and Silva ( 1984 ) reported remains possibly referable to Sterna (sensu lato) from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. The material is represented by a complete synsacrum, a partial right coracoid, a complete right humerus, the proximal end of a left ulna, and the proximal end of a right carpometacarpus. The authors noted that these remains probably represent a single species, somewhat smaller than Sterna eurygnatha (= Thalasseus acuflavidus eurygnathus ).
402. Laridae indet. 3
Mendes and Rodrigues ( 2019 , 2024 ) reported an indeterminate larid bone from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
Larinae Rafinesque
Larus Linnaeus
403. Larus sp.
Late Holocene – SC
Larus sp – Gilson and Lessa 2021: 226.
Gilson and Lessa ( 2021 ) reported 16 bones belonging to at least three individuals from the second occupation of the Rio do Meio site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
404. Larus dominicanus Lichtenstein
Holocene – SC
Larus dominicanus – Gilson and Lessa 2021: 226.
Gilson and Lessa ( 2021 ) reported seven bones belonging to at least two individuals from the second occupation of the Rio do Meio site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
Mendes and Rodrigues ( 2019 , 2024 ) reported four bones belonging to at least two individuals from the Caieira and Congonhas I (SC-LS-30) sambaquis of Laguna and Tubarão, respectively, in Santa Catarina. The middle and distal portions of a left humerus and a right femur from Congonhas I were depicted.
Rynchopinae Bonaparte
Rynchops Linnaeus
405. Rynchops niger Linnaeus
Mendes and Rodrigues ( 2024 ) reported two bones of at least one individual from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
Sterninae Vigors
Thalasseus Boie
406. Thalasseus sp.
Holocene – RJ
Thalasseus sp – Kneip et al. 1975: 105.
Thalasseus sp – Kneip 1977: 52.
Thalasseus sp – Souza Cunha et al. 1977: 147.
“ Thalasseus sp., este hoje Sterna ” – Ruschi 1979: 36.
Thalasseus sp – Mendonça de Souza and Mendonça de Souza 1981 –1982: 115.
Thalasseus sp. [in part?] – Gaspar 2003: 58.
Kneip et al. ( 1975 ) reported material from layer III of Sambaqui do Forte, in Cabo Frio, Rio de Janeiro. It consists of some bones from the wing and leg.
407. Thalasseus acuflavidus (Cabot)
Holocene – RJ
Sterna eurygnatha – Kneip et al. 1994: 51.
Kneip et al. ( 1994 ) reported three bones of at least one individual from layer II (3,960±200 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
Sphenisciformes Sharpe
Spheniscidae Bonaparte
Spheniscus Brisson
408. Spheniscus magellanicus (Forster)
Holocene – PR, RJ, RS, SC
Spheniscus magellanicus [in part] – Jacobus and Gil 1987: 116.
Spheniscus magellanicus [in part] – Jacobus et al. 1988: 465.
Spheniscus magellanicus [in part?] – Gazzaneo et al. 1989: 127.
Spheniscus magellanicus [in part] – Jacobus 1991: 74.
“Penguin bones” [in part] – Sick 1993: 119.
Spheniscus magellanicus [in part] – Rosa 1996: 159.
“Ossos de pingüins” [in part] – Sick 1997: 186.
Jacobus and Gil ( 1987 ) reported seven fragments of at least two individuals from the upper layer of square A2 of the Itapeva (RS-LN-201) site in Torres, Rio Grande do Sul. They were discovered in the first half of 1982 by the staff of the Centro de Estudos e Pesquisas Arqueológicas (CEPA) of PUCRS (Gazzaneo et al. 1989 ). The material is represented by a cervical vertebra, a complete left coracoid, the distal part of another left coracoid, the proximal part of a left humerus, a right humerus, and two distal right tibiotarsi.
Jacobus et al. ( 1988 ) reported additional material (not specifying whether this includes the seven fragments reported above) from that site, in 12 m 2 out of 80 m 2 excavated. The material comprises 22 fragments of at least five individuals, represented by the front and hindlimbs, coracoids, and synsacra, and is about 4,000 years old.
Gazzaneo et al. ( 1989 ), complementing Jacobus and Gil ( 1987 ), reported eight individuals represented by the coracoid, humerus, femur, tibiotarsus, and tarsometatarsus from the second (A3), third (A2 and B2), and fourth (A3 and B1) occupations of the site.
Rosa ( 1996 ) reported further remains of at least three individuals, from the second, third, and fourth layers of the site, excavated in the campaign’s second stage (squares C1, C2, C3, D1, D2, and D3).
Jacobus et al. ( 1988 ) reported 116 fragments (about 1,000 years old) of at least 19 individuals from the Praia das Laranjeiras II site in Balneário Camboriú, Santa Catarina. The material is represented by the left and right coracoids, left and right humeri, left and right ulnae, left and right radii, left carpometacarpus, left and right femora, left and right tibiotarsi, right tarsometatarsus, and synsacrum (Schmitz et al. 1993 ).
Schmitz and Bitencourt ( 1996b ) reported this species from the Pântano do Sul (SC-F-10) site in Ilha de Santa Catarina, Florianópolis, Santa Catarina. It makes up the bulk of the avian remains found there, with a total of 1,326 bones with known provenance within the site, including almost 200 vertebrae. Bones like the scapula, cranium, mandible, carpometacarpus, phalanges, radius, and sternum are generally completely preserved. Meanwhile, others show a high percentage of broken elements: 97 of a total of 197 coracoids are broken, and so are 83 of 133 humeri, 72 of 126 ulnae, 153 of 295 femora, 185 of 198 tibiotarsi, and 29 of 50 tarsometatarsi. Most of the fractures were the product of human action.
“pinguins” – Silva et al. 1990: 139.
Silva et al. ( 1990 ) mentioned penguin bones (despite noting that they might not have been well recognized and were not included in their archeofaunal list) from the Praia da Tapera site in Florianópolis, Santa Catarina.
Spheniscus magellanicus – Bandeira 1992: 81.
Bandeira ( 1992 ) reported this species from Sambaqui Enseada I (SC-LN-71) in São Francisco do Sul, Santa Catarina. The analyzed material is represented by 47 bones belonging to at least four individuals from the site’s first occupation and 70 bones belonging to at least seven individuals from the site’s second occupation.
“Pinguim de Magalhães” – Schmitz et al. 1992: 110.
Schmitz et al. ( 1992 ) reported this species from the Armação do Sul site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
Kneip et al. ( 1989b , 1994 ) reported four bones of at least one individual from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Kneip et al. ( 1994 ) reported a bone from layer I (3,610±190 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
Spheniscus magellanicus – Schmitz and Bitencourt 1996a: 41.
Schmitz and Bitencourt ( 1996a ) reported this species from the Praia das Laranjeiras I site in Balneário Camboriú, Santa Catarina.
“pingüim” – Schmitz and Verardi 1996: 143.
Schmitz and Verardi ( 1996 ) reported penguin remains from the Cabeçudas site in Itajaí, Santa Catarina.
Rosa ( 1995 –1996) reported a bone fragment possibly attributable to this species among the material from squares F2 and E3 of the Barra Velha (SC-IÇ-01) site in Içara, Santa Catarina.
Later, from squares E4 and W7, Rosa ( 1999 ) reported a coracoid, two humeri, a radius, and an ulna.
Spheniscus sp. [in part] – Teixeira and Rosa 2001: C00003.
Spheniscus sp. [in part] – Teixeira 2006: 21.
Spheniscus magellanicus [in part] – Rosa 2006a: 41.
Spheniscus magellanicus [in part] – Tamiozzo 2007: 289.
Spheniscus magellanicus [in part] – Tamiozzo et al. 2008: 110–111.
Spheniscus sp. [in part] – Campos 2015: 61.
Spheniscus magellanicus [in part] – Pavei 2019: 22.
Teixeira and Rosa ( 2001 ) reported 273 bones belonging to at least 46 individuals from the Rincão (SC-IÇ-06) site in Içara, Santa Catarina. The material represents at least 38 individuals according to Teixeira ( 2006 ) and is deposited in the IAP/Unisinos collection.
Tamiozzo ( 2007 ) and Tamiozzo et al. ( 2008 ) reported further material from the site. It is represented by crania (<5), mandibles (<5), vertebrae (<45), scapulae (<10), humeri (<20), ulnae (<20), radii (<10), carpometacarpi (<25), femora (<20), tibiotarsi (<20), phalanges (<5), and synsacra (<10). A total of 224 elements are mentioned in their Table 1 .
“pingüim” – Gaspar 2003: 58.
Gaspar ( 2003 ) reported penguin material (most likely this species) from the Ponta da Cabeça site in Arraial do Cabo, Rio de Janeiro.
Spheniscus magellanicus – Brentano et al. 2006: 213.
Brentano et al. ( 2006 ) reported remains from the upper layers of the RS-LC-97 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Spheniscus magellanicus – Rosa 2006d: 265.
Rosa ( 2006d ) reported a tarsometatarsus fragment from the late Holocene of the RS-LC-81 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Klokler ( 2008 ) reported this species from the Jabuticabeira II site in Jaguaruna, Santa Catarina.
Spheniscus magellanicus – Ramos Júnior 2014: 97.
Ramos Júnior ( 2014 ) reported a vertebra from Sambaqui Ilha das Pedras in Paranaguá, Paraná.
Spheniscus magellanicus [in part] – DeBlasis et al. 2014: 122.
Spheniscus magellanicus [in part] – Cardoso et al. 2014: 150–170, figs 4–5.
Spheniscus magellanicus [in part] – Cardoso et al. 2016: 1–5.
Spheniscus magellanicus – Cardoso 2018: 93, fig 21.
Spheniscus magellanicus – Cardoso et al. 2019: 71.
DeBlasis et al. ( 2014 ) reported seven bones belonging to at least one individual from level 3 of Galheta IV in Laguna, Santa Catarina, a site associated with a Jê occupation dated 1400–787 years BP (Cardoso et al. 2019 ).
Cardoso et al. ( 2014 , 2016 ) analyzed the total material collected during 2005–2007 fieldwork by GRUPEP-Arqueologia/UNISUL and MAE/USP (and thus including the material described by DeBlasis et al.) and reported the species’ remains to be collected from all the site’s five levels, in 229 of a total of 1234 analyzed archeofaunal samples, with 444 identified bones representing at least 35 individuals. Despite being apparently homogenously distributed, these elements were found in closer association with human burials 2, 4, 6, 7, and 8, which indicates a possible symbolic importance. The material is represented by the articulars (2 right), orbital fragments (4), quadrates (4 right, 6 left), coracoids (24 right, 14 left), scapulae (11 right, 10 left), sterna (2), humeri (27 right, 35 left), radii (31 right, 15 left), ulnae (18 right, 14 left), carpometacarpi (11 right, 7 left), wing phalanges (13), femora (14 right, 23 left), pedal phalanges (91), tibiotarsi (24 right, 12 left), tarsometatarsi (15 right, 8 left), synsacra (5), and caudal vertebrae (4). The limbs account for 80.6% of the total material. Of the total material, 178 bones had cutting marks, and 39 had burning traces associated with human activities.
Cardoso’s ( 2018 ) analysis of the material from the site resulted in 142 elements belonging to at least 13 individuals: five bones (belonging to at least one individual) from area A’s square 110/94, 36 bones (belonging to at least three individuals) from square 111/99, 35 bones (belonging to at least three individuals) from square 112/93, seven bones (belonging to at least one individual) from square 113/95 (the material reported by DeBlasis et al.), 54 bones (belonging to at least four individuals) from area B, and five bones (belonging to at least one individual) from the “profile” area. The material is represented by the skull (beak, frontal, articular, quadrate), sternum, ribs, synsacrum, pelvis, caudal vertebrae, pygostyle, scapula, coracoid, furcula, humerus, radius, ulna, carpometacarpus, wing phalanges, femur, patella, tibiotarsus, fibula, tarsometatarsus, and pedal phalanges.
Spheniscus magellanicus – Pavei et al. 2015: 74, 79, figs A, C in pag. 88.
Pavei et al. ( 2015 ) reported 95 bones (including humeri, femora, and tibiotarsi) representing at least 78 individuals from Sambaqui do Papagaio in Bombinhas, Santa Catarina.
Mendes and Rodrigues ( 2019 , 2022 , 2024 ) reported 34 bones belonging to at least 10 individuals from the Guaraguaçu B (Pontal do Paraná, Paraná), Caieira (Laguna, Santa Catarina), and Figueira II (Arroio do Sal, Rio Grande do Sul) sambaquis. A right coracoid from Guaraguaçu B, a right humerus and a right femur from Caieira, and the distal portion of a tibiotarsus from Figueira II were depicted.
Spheniscus magellanicus – Gilson and Lessa 2021: 226.
Gilson and Lessa ( 2021 ) reported nine bones belonging to at least one individual from the second occupation of the Rio do Meio site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
Procellariiformes Fürbringer
409. Procellariiformes indet. 1
DeBlasis et al. ( 2014 ) reported indeterminate procellariiforms from the Galheta IV site in Laguna, Santa Catarina. They associated seven bones with at least two individuals, from the site’s levels 3 (six bones) and 5 (one bone) (area A’s square 113/95 of Cardoso 2018 ).
410. Procellariiformes indet. 2
Holocene – SC
Procelariforme [sic] – Pavei et al. 2015: 74, 78, fig. B in pag. 88.
Pavei et al. ( 2015 ) associated with this order 26 bones representing at least 17 individuals from Sambaqui do Papagaio in Bombinhas, Santa Catarina.
Diomedeidae Gray
411. Diomedeidae indet.
Schmitz et al. ( 1992 ) reported dioemedeid remains from the Armação do Sul site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
Diomedea Linnaeus
412. cf. Diomedea sp.
Rosa ( 1999 ) reported an ulna comparable to this genus (sensu lato?) from square W15 of the Barra Velha (SC-IÇ-01) site in Içara, Santa Catarina.
Thalassarche Reichenbach
413. cf. Thalassarche sp.
Holocene – SC
cf. Thalassarche sp. – Pavei et al. 2015: 74, 78.
Pavei et al. ( 2015 ) reported two bone fragments comparable to this genus representing two individuals from Sambaqui do Papagaio in Bombinhas, Santa Catarina.
414. Thalassarche sp. 1
Rosa ( 2008a ) reported Thalassarche material with human-made cutting marks from the Capão Alto (RS-LN-19) site in Xangri-Lá, Rio Grande do Sul.
415. Thalassarche sp. 2
DeBlasis et al. ( 2014 ) reported this genus from the Galheta IV site in Laguna, Santa Catarina. They associated 97 bones with at least 8 individuals, from the site’s levels 1 (34 bones), 3 (44 bones), and 5 (19 bones). Some long bones were modified as tools, showing cut and polishing markings.
Cardoso ( 2018 ) analysis of the material from the site resulted in 1,020 elements belonging to at least 48 individuals: 27 bones (belonging to at least two individuals) from area A’s square 110/94, 286 bones (belonging to at least 12 individuals) from square 111/99, 260 bones (belonging to at least ten individuals) from square 112/93, 97 bones (belonging to at least eight individuals) from square 113/95 (the material reported by DeBlasis et al.), 272 bones (belonging to at least eight individuals) from area B, and 78 bones (belonging to at least eight individuals) from the “profile” area. The material is represented by the skull (beak, frontal, articular, occipital, neurocranium, quadrate), atlas, sternum, ribs, synsacrum, pelvis, scapula, coracoid, furcula, humerus, radius, ulna, carpometacarpus, wing phalanges, femur, patella, tibiotarsus, fibula, tarsometatarsus, and pedal phalanges.
416. Thalassarche sp. 3
Middle Holocene – RS
Thalassarche sp. – Mendes and Rodrigues 2024: 8, 4D.
Mendes and Rodrigues ( 2024 ) associated with this genus three bones of at least two individuals Sambaqui da Figueira II in Arroio do Sal, Rio Grande do Sul. The distal portion of a left tarsometatarsus was depicted.
417. Thalassarche cf. chlororhynchos (Gmelin)
Middle Holocene – RJ
Procellariiformes [?; in part] – Kneip et al. 1988: 48.
Procellariformes [sic] [in part] – Kneip et al. 1989a: 126.
Diomedea cf. chlororhyncus [sic] – Kneip et al. 1989b: 662.
Diomedea cf. chlororhynchos – Kneip et al. 1994: 48.
Diomadeidae (“albatroz”) [sic] [in part] – Kneip et al. 1995: 7.
Diomadeidae (“albatroz”) [sic] [in part] – Kneip et al. 1997: 19, 37.
Diomadeidae [sic] [in part] – Kneip 2001: 7.
Diomadeidae (“albatroz”) [sic] [in part] – Magalhães et al. 2001: 59.
Kneip et al. ( 1989b , 1994 ) reported a bone comparable to this species from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
418. Thalassarche cf. melanophris (Temminck) 1
Quaternary – RJ
Procellariiformes [?; in part] – Kneip et al. 1988: 48.
Procellariformes [sic] [in part] – Kneip et al. 1989a: 126.
Diomedea cf. melanophris – Kneip et al. 1989b: 662.
Diomedea cf. melanophris [in part] – Kneip et al. 1994: 48.
Diomadeidae (“albatroz”) [sic] [in part] – Kneip et al. 1995: 7.
Diomadeidae (“albatroz”) [sic] [in part] – Kneip et al. 1997: 19, 37.
Diomadeidae [sic] [in part] – Kneip 2001: 7.
Diomadeidae (“albatroz”) [sic] [in part] – Magalhães et al. 2001: 59.
Kneip et al. ( 1989b , 1994 ) reported material comparable to this species from Sambaqui da Beirada in Saquarema, Rio de Janeiro. It comprises three bones of at least one individual from the site’s layer I (3,800±190 years BP), five bones of at least three individuals from layer II (4,160±180 years BP), two bones of at least one individual from layer III (4,300±190 years BP), and two bones of at least one individual from layer IV (4,520±190 years BP).
419. Thalassarche cf. melanophris (Temminck) 2
Quaternary – RJ
Diomedea cf. melanophris [in part] – Kneip et al. 1994: 48.
Diomadeidae (“albatroz”) [sic] [in part] – Kneip et al. 1995: 7.
Diomadeidae (“albatroz”) [sic] [in part] – Kneip et al. 1997: 19, 37.
Diomadeidae [sic] [in part] – Kneip 2001: 7.
Diomadeidae (“albatroz”) [sic] [in part] – Magalhães et al. 2001: 59.
Kneip et al. ( 1994 ) reported six bones of at least one individual comparable to this species from layer II (3,960±200 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
420. Thalassarche cf. melanophris (Temminck) 3
Quaternary – SC
Diomedea sp. [in part] – Teixeira and Rosa 2001: C00003.
Diomedea sp. [in part] – Teixeira 2006: 21.
Diomedea sp. [in part] – Rosa 2006a: 41.
Thalassarche sp. [in part] – Tamiozzo 2007: 289.
Talasarche sp. [sic] [in part] – Tamiozzo et al. 2008: 110–111.
Thalassarche sp. [in part] – Campos 2015: 61.
Teixeira and Rosa ( 2001 ) reported 240 bones belonging to at least 52 individuals from the Rincão (SC-IÇ-06) site in Içara, Santa Catarina. The material represents at least 39 individuals according to Teixeira ( 2006 ) and is deposited in the IAP/Unisinos collection. The authors noted it possibly represents Thalassarche melanophris .
Tamiozzo ( 2007 ) and Tamiozzo et al. ( 2008 ) reported further material from the site. It is represented by crania (about 10), mandibles (<5), vertebrae (<20), scapulae (<5), humeri (<10), ulnae (<5), carpometacarpi (<10), femora (<5), tibiotarsi (<20), tarsometatarsi (<25), phalanges (about 40), and synsacra (about five). A total of 144 elements are mentioned in their Table 1 .
421. Thalassarche cf. melanophris (Temminck) 4
Quaternary – RS
[cf.] Diomedea melanophrys [lapsus] – Lopes et al. 2006a: 45.
[cf.] Thalassarche melanophrys [lapsus] – Lopes et al. 2006b: 648–650, 2A–C.
Thalassarche melanophrys [lapsus] – Hsiou 2007: 23.
[cf.] Thalassarche melanophrys [lapsus] – Hsiou 2009a: 150, 9B–E.
Thalassarche melanophrys [lapsus] – Lopes and Pereira 2017: 96.
cf. Thallasarche melanophris [lapsus] – Lopes et al. 2019: 200.
Fossils attributable to this species from submerged fossiliferous deposits of late Pleistocene age along the internal continental shelf of the state of Rio Grande do Sul were first described by Lopes et al. ( 2006a ) and later with further details by Lopes et al. ( 2006b ).
The material consists of the middle part of the diaphysis of a left tibiotarsus (LGP-A0002) collected at the “concheiros” region, a cervical vertebra (LGP-A0001) collected at Farolete da Verga, and another cervical vertebra (LGP-A0003) collected at Farol Sarita, all three between 1999 and 2000 during fieldwork in the coastline south of Laguna dos Patos. A third cervical vertebra (LGP-A0004) was collected near Balneário do Cassino in 2003 [32] .
The material’s good condition indicates that no significant reworking had occurred from their source sites to the beach and suggests that the submerged deposits are located near the present shoreline. In addition, remains of terrestrial mammalian megafauna, cetaceans, elasmobranchs, teleosts, crustaceans, echinoderms, and mollusks were found in the same deposits.
422. Thalassarche cf. melanophris (Temminck) 5
Ferrasso et al. ( 2013 ) reported material comparable to this species from Sambaqui Arroio Seco V (RS-LN-285) in Arroio do Sal, Rio Grande do Sul. The remains are represented by the scapula, carpometacarpus, phalanx, and tibiotarsus.
423. Thalassarche melanophris (Temminck)
Late Holocene – SC
Diomedea melanophris – Bandeira 1992: 81.
Bandeira ( 1992 ) reported this species from Sambaqui Enseada I (SC-LN-71) in São Francisco do Sul, Santa Catarina. The analyzed material is represented by five bones belonging to at least two individuals from the site’s first occupation, and 67 bones belonging to at least nine individuals from the site’s second occupation.
Procellariidae Leach
424. Procellariidae indet. 1
Carvalho ( 1984 ) reported procellariid remains among the material discovered in 1978 at the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
425. Procellariidae indet. 2
Late Holocene – RJ
Procellaridae – Lima and Silva 1984: 28.
Lima and Silva ( 1984 ) reported procellariid remains from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. The material is represented by the proximal ends of a right and a left humeri and fragments of long bone diaphyses.
426. Procellariidae indet. 3
Teixeira and Rosa ( 2001 ) reported five bones from the Rincão (SC-IÇ-06) site in Içara, Santa Catarina. The material represents at least four individuals according to Teixeira ( 2006 ) and is deposited in the IAP/Unisinos collection.
Campos ( 2015 ) cited Teixeira ( 2006 ) and Rosa ( 2006a ) and included an indeterminate larid among remains from SC-IÇ-06, but it probably refers to this record instead.
427. Procellariidae indet. 4
Holocene – SC
“Procelariforme” – Ricken et al. 2014: 116.
Ricken et al. ( 2014 ) reported a procellariiform bone from Sambaqui da Rua 13 in Bombinhas, Santa Catarina.
428. Procellariidae indet. 5
Middle Holocene – SC
Procellariidae indet. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) reported two indeterminate procellariid bones of at least one individual from the Caieira sambaqui in Laguna, Santa Catarina.
Macronectes Richmond
429. Macronectes giganteus (Gmelin)
Holocene – RJ
Macronectes giganteus – Kneip et al. 1994: 48.
Kneip et al. ( 1994 ) reported a bone from layer I (3,610±190 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
Procellaria Linnaeus
430. Procellaria aequinoctialis Linnaeus
Holocene – RJ
Procellariiformes [? in part] – Kneip et al. 1988: 48.
Procellariformes [sic] [in part] – Kneip et al. 1989a: 126.
“Procellariidae de grande porte” – Kneip et al. 1989b: 662.
“Procellariid remains” [?; in part?] – Sick 1993: 114.
Procelleria aequinoctialis [sic] [in part] – Kneip et al. 1994: 48.
“Restos de procelarídeos” [?; in part?] – Sick 1997: 178.
Kneip et al. ( 1989b ) reported a large procellariid from Sambaqui da Beirada in Saquarema, Rio de Janeiro. It was later determined as Procellaria aequinoctialis by Kneip et al. ( 1994 ). The material comprises three bones of at least one individual from the site’s layer I (3,800±190 years BP), two bones of at least one individual from layer II (4,160±180 years BP), and five bones of at least two individuals from layer III (4,300±190 years BP).
Procelleria aequinoctialis [sic] [in part] – Kneip et al. 1994: 48.
Kneip et al. ( 1994 ) reported a bone from layer I (3,610±190 years BP) and six bones of at least one individual from layer II (3,960±200 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
Ardenna Reichenbach
431. Ardenna grisea (Gmelin)
Middle Holocene – RJ
Procellariiformes [?; in part] – Kneip et al. 1988: 48.
Procellariformes [sic] [in part] – Kneip et al. 1989a: 126.
Puffinus griseus [in part] – Kneip et al. 1989b: 662.
“Procellariid remains” [?; in part?] – Sick 1993: 114.
Puffinus griseus – Kneip et al. 1994: 49.
“Restos de procelarídeos” [?; in part?] – Sick 1997: 178.
Kneip et al. ( 1989b ) reported at least one individual from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro. Later, Kneip et al. ( 1994 ) reported the material associated with this species to comprise five bones of at least two individuals.
432. Ardenna gravis (O’Reilly)
Holocene – RJ
Puffinus gravis – Kneip et al. 1994: 49.
Kneip et al. ( 1994 ) reported a bone from layer I (3,610±190 years BP) and five bones of at least two individuals from layer II (3,960±200 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
Puffinus Brisson
433. Puffinus sp. 1
Teixeira and Rosa ( 2001 ) reported 31 bones belonging to at least 14 individuals from the Rincão (SC-IÇ-06) site in Içara, Santa Catarina. The material represents at least 17 individuals according to Teixeira ( 2006 ) and is deposited in the IAP/Unisinos collection.
Tamiozzo et al. ( 2008 ) reported further material from the site, which probably belongs to the same taxon as the material reported by Teixeira ( 2006 ). It is represented by a cranium, a vertebra, a coracoid, six humeri, a radius, three metacarpi, four carpi, four femora, two tibiotarsi, a fibula, three tarsometatarsi, four phalanges, and a synsacrum, although, on their Table 1 , they mentioned a total of 28 elements.
434. Puffinus sp. 2
Middle Holocene – SC
Puffinus sp. – Mendes and Rodrigues 2024: 8, 4A.
Mendes and Rodrigues ( 2024 ) associated with this genus four bones of at least one individual from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina. The proximal and middle portions of a right humerus were depicted.
435. Puffinus puffinus (Brünnich)
Holocene – RJ
Procellariiformes [?; in part] – Kneip et al. 1988: 48.
Procellariformes [sic] [in part] – Kneip et al. 1989a: 126.
Puffinus puffinus – Kneip et al. 1989b: 662.
“Procellariid remains” [?; in part?] – Sick 1993: 114.
Puffinus puffinus – Kneip et al. 1994: 49.
“Restos de procelarídeos” [?; in part?] – Sick 1997: 178.
Kneip et al. ( 1989b , 1994 ) reported three bones of at least one individual from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Ciconiiformes Bonaparte
Ciconiidae Sundevall
436. Ciconiidae indet. 1
Quaternary – BA
Ciconiidae – Cartelle and Santos 1985: 140.
Cartelle and Santos ( 1985 ) reported a complete ciconiid skeleton among the numerous avian remains of quaternary age from Gruta dos Brejões in Morro do Chapéu, Bahia.
437. Ciconiidae indet. 2
Late Holocene – SE
Ciconidae [sic] – Queiroz and Chaix 1999: 55, 35.
Queiroz and Chaix ( 1999 ) reported ciconiid bones associated with human remains from burial 147 of the Justino site in Canindé de São Francisco, Sergipe. The material consists of two ulnae and two tibiotarsi, placed in the front of the human thorax, and is deposited in the MAX collection.
438. Ciconiidae indet. 3
Late Holocene – MS
Ciconidae [sic] – Schmitz et al. 2009: 342.
Schmitz et al. ( 2009 ) reported a ciconiid bone from level 2 of the MS-MA-16A site in Corumbá, Mato Grosso do Sul.
439. Ciconiidae indet. 4
Late Holocene – RS
Ciconiiformes – Milheira et al. 2019: 45.
Milheira et al. ( 2019 ) reported ciconiiform remains among the material from the sites PSG-02 and PSG-07 of the Pontal da Barra cerrito complex in Pelotas, Rio Grande do Sul.
Ciconia Brisson
440. Ciconia sp.
Lopes and Pereira ( 2017 ) reported an almost complete, large cervical vertebra (MCTFM-PV1090) from the fluvial deposit of the Santa Vitória Formation ( Fig. 1 .16) exposed along the right bank of Arroio Chuí in Santa Vitória do Palmar, Rio Grande do Sul. It was found isolated and disarticulated, associated with mammalian and turtle remains in sediments dated 37,900±5,080 years, and represents the first avian record for that Formation.
The specimen was initially attributed to a large anatid closely resembling Cygnus melancoryphus , but likely a distinct species. Later, Lopes et al. ( 2019 ) attributed it to Ciconia , and noted it is similar to Ciconia maguari . However, its larger dimensions suggest it could be a morphotype of that taxon or represent another extinct one, such as Ciconia lydekkeri , of which the unavailability of detailed published descriptions or specimes of cervical vertebrae for comparison prevented a more assertive attribution. The authors also discussed the role of climate change on the distribution of ciconiids and other taxa during the Quaternary in southern South America.
441. Ciconia maguari (Gmelin)
Kneip et al. ( 1989b ) reported at least one individual from layer II of Sambaqui da Beirada in Saquarema, Rio de Janeiro, but were unsure whether it belonged to Jabiru mycteria or Ciconia maguari . Later, Kneip et al. ( 1994 ) determined the material as C. maguari and reported it to comprise three bones of at least one individual from the site’s layer II (4,160±180 years BP) and six bones of at least two individuals from layer III (4,300±190 years BP).
Euxenura maguari [in part] – Kneip et al. 1994: 49.
Kneip et al. ( 1994 ) reported two bones of at least one individual from layer III (1,810±40 years BP) of Sambaqui da Pontinha in Saquarema, Rio de Janeiro.
442. † Ciconia lydekkeri (Ameghino)
Late Pleistocene – MG
Figs 24, 35B
Type locality: The assigned localities are “South America” (Lydekker 1891: 65) and “Brazilian caves” (Ameghino 1891: 445). Here, we restrict it to the Lagoa Santa region.
Etymology: lydekkeri , honoring British naturalist Richard Lydekker (1849–1915) (British Museum (Natural History)).
Palæociconia australis (Moreno) – Lydekker 1891: 64–65, 15.
Prociconia Lydekkeri nom. n. [new genus and species] – Ameghino 1891: 445.
Prociconia Lydekkeri – Lambrecht 1921: 26.
Palaeociconia australis – Lambrecht 1930: 24–25.
Palaeociconia australis (Moreno) – Lambrecht 1933: 321, 735.
Paleociconia australis [sic] – Howard 1942: 189.
Jabiru lydekkeri – Patterson and Kraglievich 1960: 8, footnote.
Prociconia lydekkeri – Brodkorb 1963: 290–291.
“Lydekker’s stork, Prociconia lydekkeri ” – HRH The Prince Philip, Duke of Edinburgh and Fisher 1970: 193.
Prociconia lydekkeri – Mones 1986: 80.
Prociconia lydekkeri – Cuello 1988: 9.
‘ Prociconia ’ lydekkeri – Haarhoff 1988: 300.
Jabiru lydekkeri – Alvarenga 1993a: 62.
Prociconia lidekkeri [lapsus] – Alvarenga 1993b: 21.
Jabiru lydekkeri – Alvarenga 1997: 21.
Ciconia lydekkeri – Agnolín 2009a: 53–58.
Ciconia lydekkeri – Nascimento and Silveira 2020: 490, 4.
Ciconia lydekkeri , holotype: proximal end of the right tarsometatarsus (NHMUK PV OR 18878), anterior (A), lateral (B), medial (C), posterior (D), proximal (E), and distal views; distal end of the left tarsometatarsus (NHMUK PV OR 18879), anterior (G), lateral (H), medial (I), posterior (J), proximal (K), and distal (L) views; referred material: distal end of the left femur (NHMUK PV OR 12878), anterior (M), medial (N), and posterior (I) views (S. Chapman; F.L. Agnolín). Scale bar: 10 mm.
Lydekker ( 1891 ) associated two fragments collected and sold by Peter Claussen to the British Museum (Natural History) with Palaeociconia australis , an Argentinean species described by Moreno ( 1889 ) from a partial tarsometatarsus from Monte Hermoso (Pliocene), Buenos Aires. The material consists of the proximal end of a right tarsometatarsus (NHMUK PV OR 18878; acquired in 1842) and the distal end of a left tarsometatarsus (NHMUK PV OR 18879).
Ameghino ( 1891 ) did not consider P. australis to be a ciconiid (it is in fact a phorusrhacid Agnolín 2009a ) from figures of the fossil later published by Moreno and Mercerat ( 1891 ) and erected the new genus and species Prociconia lydekkeri for the Brazilian material. He also provisionally classified under this name the bones of a large bird from the Pampean Formation of Buenos Aires mentioned by Burmeister.
Lambrecht ( 1930 , 1933 ), who considered this taxon one of the most problematic forms of South American fossil birds, used the name Palaeociconia australis and treated Prociconia lydekkeri as a synonym. He had no access to Moreno’s description of P. australis , and, therefore, believed he described the tarsometatarsi from Lagoa Santa in the British Museum, with Lydekker only depicting one of them. He noted, however, that this material is not identical to the one later depicted by Moreno and Mercerat, which he thought should be classified as a phorusrhacid.
Howard ( 1942 ), in her review of American fossil storks, expressed regret that she was not able to examine the P. australis material in hands, as opposed to the North American fossil ciconiid material. The two specimens originally associated with the taxon were removed from the British Museum of Natural History and stored in safer quarters due to the war, being unavailable for study when Henry Anson Wylde of the Los Angeles Museum’s Department of Paleontology visited the institution in late 1939. Based on the drawings published by Lydekker, she commented that the tarsometatarsus does not resemble the North American ones but noted that old line drawings are not always to be relied on for accuracy.
Patterson and Kraglievich ( 1960 ) followed Ameghino and considered the name given by Moreno in 1889 a nomen nudum but associated it with the genus Jabiru after an examination of the type. Brodkorb ( 1963 ) used Prociconia lydekkeri and found no evidence to place the species in the genus Jabiru .
Harrison ( 1975 ) proposed that Lydekker’s publication constituted validity for the name Palaeociconia australis attributed to the material of the Lagoa Santa region, with Prociconia lydekkeri being a synonym. Harrison did not seem to have consulted Moreno ( 1889 ) (although he included the study in his references) and, like Lambrecht, understood he described the material from Brazil. According to him, this name given by Moreno in 1889 is a nomen nudum, and the date of publication of Lydekker’s study precedes that of Moreno and Mercerat (April 25 or “before May 2” versus “May to August 5”). Furthermore, he mentioned that Brodkorb also supported this after reviewing the data. Harrison also referred to this species the distal end of a left femur (NHMUK PV OR 12878 [33] ) from the same deposits and collection in the British Museum and suggested affinity to the genus Mycteria . This bone and the two others mentioned above are possibly referable to the same individual (Agnolín 2009a ).
Haarhoff ( 1988 ) termed it as “ Prociconia ” lydekkeri and, through personal communication with Olson, noted it required revision. This was followed by Louchart et al. ( 2005 ), who further noted that its status as a ciconiid was not certain.
Agnolín ( 2009a ) transferred the species to the genus Ciconia , considering the description of Ameghino as valid and creating the combination Ciconia lydekkeri . When comparing other fossil storks, he associated with C. lydekkeri the distal end of a right humerus from the late Pleistocene of Luján Formation in Monte Hermoso, Buenos Aires. He also synonymized Ciconia maltha to this species—a comparison already suggested by Brodkorb ( 1963 ), originally described in 1910 from a left tarsometatarsus from the late Pleistocene of Rancho La Brea in California, USA. After Miller’s description, several other specimens originally associated with C. maltha were found, including complete skeletons, with a high degree of variation, in the USA in Idaho (late Pliocene, middle and late Pleistocene), California (late Pleistocene), and Florida (late Pleistocene), and isolated fragments in Cuba (late Pleistocene of Cienfuegos province) and Bolivia (early–middle Pleistocene of Tarija) (Brodkorb 1963 , Feduccia 1967 , Agnolín 2006b ).
The synonymization of the Rancho La Brea stork with the bird of Lagoa Santa allows a clearer picture of this component of the Brazilian prehistoric avifauna. Agnolín (2006b) suggested, when describing the material collected in Bolivia, that, given the age of the fossils, it is likely that the stork came from North to South America during the Great American Biotic Interchange.
Jabiru Hellmayr
443. cf. Jabiru mycteria (Lichtenstein)
Late Holocene – PA
“tuyuyú” – Meggers and Evans 1957: 273.
Meggers and Evans ( 1957 ) mentioned four parts of bird skulls, “including one from a large species like the tuyuyú ”, besides numerous postcranial fragments and small rodent skulls, from a funerary jar with human remains found in the Guajará mound in Monte Carmelo (PA-JO-14), a Marajoara phase site in Ilha do Marajó, Pará. Both human and animal bones had been painted red.
444. Jabiru mycteria (Lichtenstein)
Kneip et al. ( 1994 ) reported three bones of at least one individual from layer II (3,960±200 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro.
Mycteria Linnaeus
445. Mycteria cf. americana Linnaeus
Late Pleistocene – RS
Ciconiidae indet. – Ribeiro et al. 1995: 107.
“ave da família Coconiidae” [sic] – Oliveira et al. 1997: 75.
cf. Ciconia sp. – Oliveira 1999: 67.
Ciconiformes [sic] – Hsiou 2006: 65.
“Ciconiiformes indeterminado” – Hsiou 2007: 23.
Ciconiiformes indet. – Kerber and Oliveira 2008a: 50, 62.
Mycteria cf. M. americana – Hsiou 2009a: 145–146, 4.
“ave ciconiforme” – Hsiou 2009b: 335–337, figs 15–16.
Mycteria cf. M. americana – Kerber et al. 2011: 204.
Mycteria cf. M. americana – Kerber et al. 2014: 254.
Ciconiidae – Lopes and Pereira 2017: 96.
cf. Mycteria americana – Lopes et al. 2019: 200.
Ribeiro et al. ( 1995 ) reported this first late Pleistocene avian record for Rio Grande do Sul. It consists of a fragment of tarsometatarsus (MCN-PV 1845) lacking the proximal end found in the Barranca Grande locality of the Touro Passo Formation in Uruguaiana (Kerber and Oliveira 2008a ).
The fragment was first determined as close to the ciconiids by Ribeiro et al. ( 1995 ) and attributed to cf. Ciconia sp. by Oliveira ( 1999 ). According to Hsiou ( 2007 , 2009a ), it is quite similar to Mycteria americana , but, due to the poor state of conservation and the absence of more skeleton elements for comparison, a definitive determination was not possible.
In the same Formation, several species of mammals were also found, including large ones, and reptiles, fishes, mollusks, and phytoliths (Kerber and Oliveira 2008a , Kerber et al. 2011 , 2014 ).
446. Mycteria americana Linnaeus
Late Holocene – SP?/PR?
Mycteria americana – Emperaire and Laming 1956: 149.
Emperaire and Laming ( 1956 ), while describing sambaquis from the southern coast of Brazil (São Paulo and Paraná), mentioned this species among rare avian remains from the shell levels.
Suliformes Sharpe
Fregatidae Degland & Gerbe
447. Fregatidae indet.
Olson ( 1981 ) reported well-mineralized seabird bones as frequent during the 1973 collection on the dunes of the Santo Antônio peninsula in Fernando de Noronha. The proximal end of a humerus appears to belong to a small species the size of Fregata ariel , which in the Atlantic Ocean is now found only on Trindade Island, although Olson ( 1975 ) reported subfossils of similar size from Saint Helena Island. A fragmented bone from the indurated sandstone caves east of the dunes was also reported by Carleton and Olson ( 1999 ). Olson (personal communication, 2016) further noted that the frigate and booby (see Sulidae indet.) material is mostly very fragmentary, and much of it may not be identifiable beyond the genus.
Fregata Lacépède
448. Fregata magnificens Mathews
Late Holocene – RJ
Fregata magnificens – Lima and Silva 1984: 29.
Lima and Silva ( 1984 ) reported remains from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. The material includes at least two individuals and is represented by two C-9 cervical vertebrae, the cranial end of a right coracoid, the cranial extremity of a right coracoid, the cranial half of a right coracoid, the proximal end of a right humerus, the distal end of a right ulna, part of the distal end of the diaphysis of a left ulna, fragment of the distal end of a left ulna, and a distal third of a left ulna.
“Fragata” – Gaspar 2003: 58.
Gaspar ( 2003 ) reported fregatid material (most likely this species) from the Ponta da Cabeça site in Arraial do Cabo, Rio de Janeiro.
Sulidae Reichenbach
449. Sulidae indet.
Holocene – PE
“Boobies” – Olson 1981: 483.
Olson ( 1981 ) reported well-mineralized, fragmentary sulid bones collected in 1973 in the dunes of the Santo Antônio peninsula in Fernando de Noronha.
Sula Brisson
450. Sula sp.
Kneip et al. ( 1975 ) reported material associated with this genus from layer III of Sambaqui do Forte, in Cabo Frio, Rio de Janeiro. It consists of a distal portion of a femur and a fragment of carpometacarpus of a bird more robust than the living Sula species.
451. Sula leucogaster (Boddaert)
Holocene – RJ, SC
Sula leucogaster leucogaster – Lima and Silva 1984: 28.
Lima and Silva ( 1984 ) reported remains from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. The material includes at least two to three individuals and is represented by cervical vertebrae (C-7, C-8, C-11, and C-12), fragments of the pelvis, an almost complete synsacrum, an incomplete right coracoid, the proximal end of a left humerus, fragments of the diaphysis of the humerus, and two fragments of the diaphysis of the ulna.
Pelicaniformes [sic] [in part] – Kneip et al. 1989a: 126.
Sula leucogaster – Kneip et al. 1989b: 662.
Sula leucogaster – Kneip et al. 1994: 49.
Sulidae (“atobá”) – Kneip et al. 1995: 7.
Sulidae (“atobá”) – Kneip et al. 1997: 19, 37.
Sulidae – Kneip 2001: 7.
Sulidae (“atobá”) – Magalhães et al. 2001: 59.
Kneip et al. ( 1989b , 1994 ) reported two bones of at least one individual from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Sula leucogaster – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) reported two bones of at least one individual from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
Anhingidae Reichenbach
452. † Anhingidae indet. 1
Souza-Filho and Guilherme ( 2015 ) mentioned anhingid material from the Talismã locality of Solimões Formation on the right bank of the Purus River, Amazonas, without developing further. Muniz et al. ( 2018 ) reported an indeterminate avian tarsometatarsus fragment collected during the 2015–2016 UFAC and USP campaign at the site. Bissaro-Júnior et al. ( 2019 ) mentioned bird material from the Talismã locality, though they cited Alvarenga and Guilherme ( 2003 ) as the source.
453. † Anhingidae indet. 2
Late Miocene – AC
Anhingidae [in part] – Hsiou et al. 2022b: 15.
Hsiou et al. ( 2022b ) preliminarily reported anhingid cranial and post-cranial elements among material collected during the July 2022 expedition carried out by the staff of USP, UFAC, and ICMBio in the Upper Acre River. Post-cranial remains were later described by Guilherme et al. ( 2023 ), without further mention of cranial remains.
Anhinga Brisson
454. † Anhinga sp.
Bandeira et al. ( 2015 ) reported the distal end of a right tarsometatarsus (MCT.R.1297) from the Cachoeira do Bandeira locality and attributed it to an indeterminate Anhinga .
455. Anhinga anhinga (Linnaeus)
Kneip et al. ( 1994 ) reported a bone from layer I (younger than 1,790±50 years BP) of Sambaqui da Pontinha in Saquarema, Rio de Janeiro.
456. †cf. Anhinga minuta Alvarenga & Guilherme
Bandeira et al. ( 2015 ) reported a fragmented right humerus (MCT.R.1168) from the Cachoeira do Bandeira locality and referred to it as Anhinga cf. A. minuta .
Anhinga sp., distal end of the right tarsometarsus (MCT.R.1297) in dorsal (A), lateral (B), and ventral (C) views; cf. Anhinga minuta: right humerus (MCT.R.1168) in cranial (D) and caudal (E) views (A-C: K.L.N. Bandeira; D–E: R.C. Silva). Scaler bar: 5 mm.
457. † Anhinga minuta Alvarenga & Guilherme
Late Miocene – AC
Fig. 34 B
Type locality: Sítio Cachoeira do Bandeira, between the Brazilian cities of Brasiléia and Assis Brasil, on the left side of the Rio Acre, Acre State, Brazil (approx. 69° 20’ W, 11° 26’ S).
Etymology: minuta , Latin for “small”.
Anhinga sp. [?; in part?] – Rancy et al. 1989: 249.
“Fósiles de ánades” [?; in part?] – Wall et al. 1991: 397.
“Anhingidae extinto” [?; in part?] – Alvarenga 1992: 254.
Anhinga sp. [?; in part?] – Bocquentin and Silva 1998: 154.
Anhinga nov. sp. [?; in part?] – Melo et al. 1998: C.4-048.
Anhinga minuta , sp. nov. [new species] – Alvarenga and Guilherme 2003: 617–618, figs 7A, 8A, 9D, 10A.
Anhinga minuta – Alvarenga and Höfling 2011: 127.
Anhinga minuta – Diederle 2015: 100–101, pl. 1.
Anhingidae [in part] – Souza-Filho and Guilherme 2015: 152.
Anhinga minuta – Hsiou et al. 2022a: 40.
Alvarenga and Guilherme ( 2003 ) described this new species from fossils of the Solimões Formation discovered at the Acre Conglomerate Member of the Cachoeira do Bandeira site (LACM 5158; Negri et al. 2010 ). It is the smallest known anhingid.
The material consists of an almost complete left tibiotarsus (UFAC-4720, holotype) and an almost complete left humerus (UFAC-4719), which show signs of advanced ontogenetic states (Diederle 2015 ).
An estimated height of about 50 cm was mentioned by Pivetta ( 2003 ). Diederle ( 2015 ) estimated its body mass at 0.9 kg and wingspan at 91.3 cm, and later (Diederle 2016 ) at 729 g and 0.958 m, respectively. He used these parameters and others (wing area and load), along with the reconstruction of its musculature, to infer the capacity of flapping flight, alternated with moments of thermal soaring, and fast flights in forested areas. The bird would have been able to dive efficiently, climb the vegetation, and take off from the water to outwit predators quickly. Their food would consist mainly of small fishes, complemented with invertebrates, amphibians, and small reptiles, and their nesting area would be the same as in the living species, on trees.
Noriega ( 1995 ) reported six isolated humeri in various states of preservation from the “Mesopotamiense”, Ituzangó Formation (late Miocene) in the province of Entre Ríos, Argentina, and noted that perhaps they would represent a new species of probably flightless Anhinga . Noriega and Agnolín ( 2008 ) observed similarities between this material and the referred humerus of Anhinga minuta , but, since they are similar in size to the living Anhinga anhinga , they remarked that additional and more complete material is necessary for a better systematic association of the Argentinean material (which they determined as cf. Anhinga minuta ) and to increase the knowledge about A. minuta . Cenizo and Agnolín ( 2010 ), in a preliminary reanalysis of this material, found enough differences not to group it with A. minuta , and preferred to provisionally refer to it as Anhingidae gen. et sp. indet., as proposed by Noriega ( 1995 ).
A. minuta – Guilherme et al. 2023: 10–11, 5.
Guilherme et al. ( 2023 ) associated with this species a relatively well-preserved nineteenth cervical vertebra (UFAC-7296) from the Patos locality at the Brazil/Peru border.
†Macranhinga Noriega
458. † Macranhinga sp. 1
Late Miocene – AC
“undescribed gigantic anhinga from the Miocene of Brazil” [?; in part?] – Campbell (in Mourer-Chauviré 1989 ): 7.
Anhinga sp. [?; in part?] – Rancy et al. 1989: 249.
“Fósiles de ánades” [?; in part?] – Wall et al. 1991: 397.
“Anhingidae extinto” [?; in part?] – Alvarenga 1992: 254.
Macranhinga [?; in part?] – Alvarenga 1993b: 21.
Anhinga sp. [?; in part?] – Bocquentin and Silva 1998: 154.
Anhinga nov. sp. [?; in part?] – Melo et al. 1998: C.4-048.
Anhinga cf. grandis – Alvarenga and Guilherme 2003: 618, 9E.
Anhinga grandis [in part?] – Alvarenga and Höfling 2011: 127.
Macranhinga sp. [in part] – Diederle 2015: 131–132, pl. 9 (9–11).
Anhingidae [in part] – Souza-Filho and Guilherme 2015: 152.
Anhinga cf. grandis - Hsiou et al. 2022a: 40.
Alvarenga and Guilherme reported the well-preserved distal end and shaft of a right humerus (UFAC-4721) from the Solimões Formation of the Cachoeira do Bandeira site (LACM 5158; Alvarenga and Guilherme 2003 , Negri et al. 2010 ).
It is larger than the specimen they attributed to Anhinga cf. fraileyi (see below) and matches in size and almost completely in morphology with a replica of the Anhinga grandis holotype, a species described from the late Miocene of Nebraska (USA) (Martin and Mengel 1975 ), with records in Florida (Becker 1987 ) and a disputed one from the middle Miocene of Colombia (Rasmussen and Kay 1992 ; referred to as Anhingidae gen. et sp. indet. by Cenizo and Agnolín 2010 ). Diederle ( 2015 ) reexamined the humerus and associated it with the genus Macranhinga .
459. † Macranhinga sp. 2
Late Miocene – AC
“nova espécie do gênero Anhinga” – Silva et al. 1997: 86.
“nova espécie de Anhinga ” – Bocquentin and Janoo 1997: 111.
Anhinga sp. [?; in part?] – Bocquentin and Silva 1998: 154.
Anhinga sp. nov. – Kellner 1998: 654, 659.
Anhinga nov. sp. [?; in part?] – Melo et al. 1998: C.4-048.
Anhinga – Kellner and Campos 1999: 247.
Anhinga cf. fraileyi – Alvarenga and Guilherme 2003: 618, figs 9C, 10B.
Anhinga fraileyi – Bocquentin and Melo 2006: 188.
Macranhinga fraileyi [in part] – Cenizo and Agnolín 2010: 498.
M. sp. cf. M. fraileyi – Cenizo and Agnolín 2010: 506.
Anhinga fraileyi [in part?] – Alvarenga and Höfling 2011: 127.
Macranhinga sp. [in part] – Diederle 2015: 131–132, pl. 9 (14–15).
Anhingidae [in part] – Souza-Filho and Guilherme 2015: 152.
Macranhinga paranensis - Hsiou et al. 2022a: 40.
Alvarenga and Guilherme ( 2003 ) reported a left humerus lacking only part of the distal end (UFAC-4562) from the Solimões Formation of the Cachoeira do Bandeira site (LACM 5158; Negri et al. 2010 ). It is most likely the same [34] found in 1996 in a very compact conglomerate on the left bank of the Acre River mentioned by Silva et al. ( 1997 ) and Bocquentin and Janoo ( 1997 ) as representing a new species of Anhinga .
The specimen has a size and aspect consistent with a humerus referred to as Anhinga fraileyi by Campbell ( 1996 ). However, Alvarenga and Guilherme noted that the attribution of any of these humeri to the same species as the A. fraileyi holotype (a tarsometatarsus) is not recommended due to the number of anhingid species present in this Formation.
Campbell ( 1996 ) described the large A. frailey (represented by the tarsometatarsus, humerus, ulna, tibiotarsus, and cervical vertebrae) from the late Miocene of neighbouring Peru, preferring not to erect a new genus. Noriega and Alvarenga ( 2002 ) suggested that A. fraileyi might belong to the genus Macranhinga , and preferred to keep its current status until new, more preserved fossils are discovered. This possibility was also reinforced by Noriega and Agnolín ( 2008 ), and Cenizo and Agnolín ( 2010 ) mentioned it as Macranhinga fraileyi , including the Brazilian material. Diederle ( 2015 , 2017 ) reexamined the A. fraileyi material and suggested that it is a junior synonym of Macranhinga paranensis (and part of the associated elements as an indeterminate anhingid), which expands its geographic distribution significantly. As for the Brazilian material, he associated it with the genus Macranhinga (Diederle 2015 ).
460. † Macranhinga sp. 3
Guilherme et al. ( 2023 ) associated with this genus a pelvic girdle fragment and the synsacrum (UFAC-6990) from the surface of the Acre Conglomerate in the Patos locality at the Brazil/Peru border. The mass of the individual to which this material belonged was estimated at 6 kg, similar to the estimated average mass for Macranhinga paranensis .
In the first assessment of the phylogenetic position of the specimen, it was recovered as a stem Anhingidae outside the monophyletic groups formed by Anhinga and Macranhinga + Giganhinga . Although its position must be considered with caution due to limited osteological information and statistical support, this result favors a position as an anhingid distinct from M. paranensis .
Diederle ( 2017 ) suggested that Anhinga fraileyi is synonymous with M. paranensis . However, the holotype right tarsometatarsus (LACM 135356) of A. fraileyi is slightly different from those of M. paranensis . Guilherme et al. noted that this tarsometatarsus was likely of an individual of intermediate size between the extant anhingas and the Macranhinga species, similar to the pattern they observed in the UFAC-6990 specimen. Considering the differences between the LACM 135356 and UFAC-6990 from the homologs of M. paranensis , they noted that Macranhinga fraileyi may be a valid taxon (sensu Cenizo and Agnolín 2010 ).
Guilherme et al. ( 2023 ) associated with a robust form of this genus a left femur lacking the distal end (UFAC-6993) from the same location in the Patos locality at the Brazil/Peru border as the UFAC-6991 femur. An average estimated mass of 8.9 kg was attributed to the individual to which the specimen belonged.
This femur and the UFAC-6991 specimen were found in situ and the same location as the UFAC-6990 pelvic girdle. Albeit neither fits it perfectly, the UFAC-6993 specimen is overall much closer to it. The authors found it unlikely that the two femora, with distinct mass estimates, represent individuals of the same population and noted it is possible to state that UFAC-6993 and the UFAC-6990 belong to the same taxon, but not to the same individual.
461. † Macranhinga sp. 4
Guilherme et al. ( 2023 ) associated with this genus the anterior portion of a synsacrum (UFAC-5086) from the Patos locality at the Brazil/Peru border. It was found 20 years earlier in a curve downstream from that site in the same locality described by Kay and Cozzuol ( 2006 ) and Latrubesse et al. ( 2010 ).
462. † Macranhinga sp. 5
Guilherme et al. ( 2023 ) associated with a gracile form of this genus an almost complete left femur (UFAC-6991) from the Patos locality at the Brazil/Peru border. An average estimated mass of 3.9 kg was attributed to the individual to which the specimen belonged, which is compatible with that estimated for Anhinga fraileyi . The authors considered unlikely that this specimen belongs to Anhinga grandis , unknown by the femur but with an estimated mass based on its holotype humerus about half that of UFAC-6991.
On the A. fraileyi material other than the holotype tarsometatarsus (see above) described by Campbell ( 1996 ) and reexamined by Diederle ( 2017 ), Guilherme et al. noted they might be associated with the UFAC-6991 femur in the same taxon, but only with the discovery of new material these questions can be fully resolved.
463. † Macranhinga sp. 6
Guilherme et al. ( 2023 ) associated with this genus a complete fourth cervical vertebra (UFAC-6989) from the Patos locality at the Brazil/Peru border. It may belong to Macranhinga fraileyi (sensu Cenizo and Agnolín 2010 ) or a species of the genus not yet described.
The similarities between this specimen and the UFAC-6992 vertebra, and those attributed to Macranhinga paranensis , and the differences observed when compared with the Anhinga vertebrae, as well as those found in the UFAC-6990 pelvic girdle, are indicators that the valid taxon for the Acre Conglomerate member may be Macranhinga fraileyi , as suggested by Cenizo and Agnolín ( 2010 ), instead of Anhinga fraileyi .
464. † Macranhinga sp. 7
Guilherme et al. ( 2023 ) associated with this genus a fifteenth cervical vertebra (UFAC-6992) from the Patos locality at the Brazil/Peru border. Like the specimen above, it may belong to M. fraileyi or another undescribed species.
465. † Macranhinga ranzii Alvarenga & Guilherme
Late Miocene – AC, AM
Fig. 34 C
Type locality: Sítio Niterói, about 20 km south of the city of Rio Branco, right bank of Acre river, State of Acre, Brazil (ca. 68° W, 10° 30’ S).
Etymology: Macranhinga , with the Greek makros, “large anhinga”, and ranzii , honoring Alceu Ranzi (1949–), for his dedication to the paleontology in the state of Acre.
“undescribed gigantic anhinga from the Miocene of Brazil” [?; in part?] – Campbell (in Mourer-Chauviré 1989 ): 7.
Anhinga sp. [?; in part?] – Rancy et al. 1989: 249.
“Fósiles de ánades” [?; in part?] – Wall et al. 1991: 397.
“Anhingidae extinto” [?; in part?] – Alvarenga 1992: 254.
“dos formas muy grandes de Brasil (K. E. Campbell, com. pers.) y Chile (Alvarenga, 1992 )” [?; in part] – Noriega 1992: 218.
Macranhinga [in part?] – Alvarenga 1997: 122.
Anhinga sp. [?; in part?] – Latrubesse et al. 1997: 112.
Anhinga sp. [?; in part?] – Bocquentin and Silva 1998: 154.
Anhinga sp. [?; in part?] – Carvalho et al. 2000: 1.
Macranhinga ranzii , sp. nov. [new species] – Alvarenga and Guilherme 2003: 614–616, 2A, 2C, 3A, 4A, 5A, 6A.
Macranhinga ranzi [sic] – Alvarenga and Höfling 2011: 127.
Macranhinga ranzii [in part] – Diederle 2015: 113–114, pl. 4.
Anhingidae indet [in part] – Diederle 2015: 122–126, pl. 7 (3–6, 11–12, 21–22).
?Anhingidae [in part] – Diederle 2015: 130, pl. 8.
Anhingidae – Souza-Filho and Guilherme 2015: 152.
Macranhinga ranzii – Hsiou et al. 2022a: 40, 2.5.
Alvarenga and Guilherme ( 2003 ) described this large new species from fossils of the Solimões Formation of the Niterói site (LACM 5954; Negri et al. 2010 ; not at the conglomerate level; Guilherme et al. 2023 ).
The material consists of a left femur lacking the distal end (UFAC-3640, holotype), a deformed right femur lacking the distal end (UFAC-4034, paratype), a complete right femur (UFAC-4860, paratype), the distal end of a right tarsometatarsus lacking the lateral trochlea (UFAC-3523), and four vertebrae representing the ninth (UFAC-4108), the fourteenth (UFAC-2212), the fifteenth (UFAC-3522), and the twentieth (UFAC-2235) cervical. The association of the referred material was questioned by Diederle ( 2015 ), who considered the vertebrae to be an indeterminate anhingid and the tarsometatarsus as of doubtful attribution to the Anhingidae.
This taxon received the title of largest known anhinga upon its description, but it currently belongs to cf. Giganhinga , from the late Miocene of Argentina (Areta et al. 2007 , Mayr 2016 ). In any case, the fossils indicate a very large anhingid, with an estimated weight of 7.8 kg (Areta et al. 2007 ) or 8.8 kg (Diederle 2015 ), and a size 20% to 25% larger than the giant Macranhinga paranensis (Alvarenga and Guilherme 2003 ; or 34% larger in mass according to Diederle 2015 ), genus and species described from the “Mesopotamiense”, Ituzaingó Formation (late Miocene) of the province of Entre Ríos, Argentina (Noriega 1992 ). A height estimate of 1.5 m was mentioned by Pivetta ( 2003 ). A third, smaller species, Macranhinga ameghinoi , was described from the middle Miocene of the province of Río Negro (Diederle and Agnolín 2017 ), and remains attributed to this genus were reported from the early–middle Miocene of the province of Santa Cruz, Argentina (Cenizo and Agnolín 2010 ).
Diederle ( 2015 ) suggested that its large size allowed prolonged dives in greater depths, with stronger strokes (but not more than the phalacrocoracids), and that it captured larger prey than the living anhingas do, competing for these resources with gharials or cetaceans. The bird would have inhabited both the upper and lower parts of large rivers basins and possibly nested in the ground on small islands or islets, safe from predators.
Noriega and Agnolín ( 2008 ) associated with M. ranzii the proximal end of a left femur (MACN PV 14371) from the Ituzangó Formation, being the first record of this species outside Brazil.
Diederle and Agnolín ( 2017 ; see also Agnolín 2016b ) suggested a close relationship between Macranhinga , Meganhinga (with Meganhinga chilensis of the early Miocene of Chile [Alvarenga 1995a ]) and Giganhinga (with Giganhinga kiyuensis of the Pliocene–Pleistocene of Uruguay; Rinderknecht and Noriega 2002 ). Guilherme et al. ( 2020 ) found two equally probable evolutionary hypotheses, none with a monophyletic Macranhinga , as G. kiyuensis was recovered as a sister species of M. ranzii . In one of these scenarios, M. chilensis was recovered as the sister species of the Macranhinga + Giganhinga clade.
Guilherme et al. ( 2020 ) tentatively attributed to this species new material from the Cajueiro locality in Boca do Acre, Amazonas. They are likely the two indeterminate avian specimens previously reported by Loboda et al. ( 2019 ) for that locality.
It consists of a virtually complete pelvic girdle articulated with the synsacrum (UFAC-6471), with a surprisingly well-preserved surface. These elements were unknown for this species. The fossil was recovered in two parts, with the cranial portion free from the rock over the sediments and the other half in situ, partially exposed within the rock, and were then glued together in the laboratory.
A mean weight of 8.6 kg was inferred for the species with this new material, making it the second heaviest anhingid known from South America, after G. kiyuensis with 25.7 kg. Behavioral information was also inferred, with morphological and myological data indicating the taxon was a proficient swimmer and diver.
Phalacrocoracidae Reichenbach
466. cf. Phalacrocoracidae
Late Holocene – PR
“un fragment d’os creux de jeune oiseau, peut-être de cormoran” – Emperaire and Laming 1956: 120.
Emperaire and Laming ( 1956 ) reported a hollow bone fragment of a young bird possibly belonging to a cormorant from human burial VI of the Ilha do Rato sambaqui in Guaratuba, Paraná.
467. † Phalacrocoracidae indet.
Late Oligocene/Early Miocene – SP
Phalacrocoracidae – Alvarenga (in Castro et al. 1988b ): 2360.
Phalacrocoracidae – Castro et al. 1988b: 2362.
Phalacrocoracidae – Alvarenga 1993a: 63.
Phalacrocoracidae – Alvarenga 1993b: 24.
Phalacrocoracidae – Alvarenga 1997: 123.
Phalacrocoracidae? gen. sp. – Carmo et al. 2024: 5.
Alvarenga (in Castro et al. 1988b , Alvarenga 1993b , 1997 ) mentioned poorly preserved, undescribed phalacrocoracid remains (a skull) from the Tremembé Formation that await the discovery of new specimens for further study.
Nannopterum Sharpe
468. Nannopterum sp.
Rosa ( 2004 ) reported two bones of at least one individual from the Paranaíba phase square 14H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
469. Nannopterum brasilianum (Gmelin)
Quaternary – MG, MS, RJ, RS, SC
Phalacrocorax brasilianus – Winge 1887: 31.
Phalacrocorax brasilianus – Goeldi 1894: 603.
Phalacrocorax brasilianus – Lambrecht 1933: 733.
Phalacrocorax olivaceus – Brodkorb 1963: 255.
Phalacrocorax olivaceus [ Phalacrocorax brasilianus ] – Mones 1986: 78.
Phalacrocorax olivaceus – Cuello 1988: 30.
Nannopterum brasilianus – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported part of a pelvis from Lapa da Escrivânia V and the distal end of a humerus and another partial pelvis from Lapa da Lagoa do Sumidouro. He noted that the two pelves differ somewhat from each other and a recent specimen, most likely due to individual variations.
Pelicaniformes [sic] [in part] – Kneip et al. 1989a: 126.
Phalacrocorax olivaceus – Kneip et al. 1989b: 662.
Phalacrocorax olivaceus – Kneip et al. 1994: 49.
Phalacrocoracidae (“biguá, biguatinga”) [in part] – Kneip et al. 1995: 7.
Phalacrocoracidae (“biguá, biguatinga”) [in part] – Kneip et al. 1997: 19, 37.
Phalacrocoracidae – Kneip 2001: 7.
Phalacrocoracidae (“biguá, biguatinga”) [in part] – Magalhães et al. 2001: 59.
Kneip et al. ( 1989b , 1994 ) reported five bones of at least two individuals from layer I (3,800±190 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Jacobus ( 2004 ) reported remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
Palacrocorax brasilianus [sic] – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported remains from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Phalacrocorax brasilianus – Schmitz et al. 2009: 342.
Schmitz et al. ( 2009 ) reported a bone from level 3 of the MS-MA-16A site in Corumbá, Mato Grosso do Sul.
Rosa ( 2010 ) reported a bone from the Guarani site RS-C-71 in Porto Alegre, Rio Grande do Sul.
Phalacrocorax brasilianus – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
Nannopterum brasilianus – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) reported a bone from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
Pelecaniformes Sharpe
Ardeidae Leach
470. cf. Ardeidae
Middle Holocene – SP
“uma tíbia de Ciconiforme, possivelmente de um Ardeidae” [sic] – Garcia 1972: 51.
Garcia ( 1972 ) reported a possible ardeid tibiotarsus from the Piaçaguera site in Cubatão, São Paulo. It is deposited in the MAE-USP collection.
471. † Ardeidae indet. 1
Castro et al. ( 1988b ) mentioned Ardeidae among the paleofauna of the Tremembé Formation. However, the summary of bird records from the Formation provided in the same publication, derived from the literature and information provided by Herculano Alvarenga, does not mention this family.
472. Ardeidae indet. 2 (spp.?)
Late Holocene – SP
“Ardeideos” – Figuti 1993: 74.
Figuti ( 1993 ) reported ardeid remains from the COSIPA sambaquis 1 and 4 in Ilha do Casqueirinho, Cubatão, São Paulo.
473. Ardeidae indet. 3
Holocene – RS
Ardeidae – Rosa 2006e: 344.
Rosa ( 2006e ) associated with this family a fragment of maxilla from the Banhado do Colégio site in Camaquã, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
474. Ardeidae indet. 4 (spp.)
Kokler ( 2008 ) reported two different ardeid species from the Jabuticabeira II site in Jaguaruna, Santa Catarina.
Ixobrychus Billberg
475. Ixobrychus exilis (Gmelin)
Quaternary – MG
Ardetta erythromelas et aff. – Winge 1887: 30–31.
Ardea ( Ardetta ) erythromelas – Goeldi 1894: 559.
Ardetta erythromelas – Lambrecht 1933: 734.
Ixobrychus exilis – Brodkorb 1963: 285.
Ixobrychus exilis [ Ardetta erythromelas ] – Mones 1986: 80.
Ixobrychus exilis – Cuello 1988: 32.
Ixobrychus exilis erythromelas – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported several elements from Lapa da Escrivânia V that he associated with the species or are close to it. He had bones from skins of Ixobrychus exilis and Ixobrychus erythromelas (= Ixobrychus exilis erythromelas ) available for comparison. There are two tarsometatarsi, one that corresponds well with that of I. e. erythromelas but only slightly more robust, and another that is somewhat smaller and markedly different; a first phalanx of the left hallux, slightly shorter than in I. e. erythromelas , but with the proximal end a little more robust; two tibiotarsi, one of which is slightly smaller and divergent than that of I. e. erythromelas ; an incomplete humerus, but which corresponds well to that of I. e. erythromelas ; two cervical vertebrae, much smaller than in Butorides striata , most likely from Ixobrychus ; a coracoid, similar to that of Butorides , but smaller and relatively longer and slender; and two femora, one longer and thinner and different in several aspects from that of B. striata , and the proximal end of another, slightly smaller than in B. striata , which may belong to another species than the other bones, perhaps B. striata itself.
Butorides Blyth
476. Butorides striata (Linnaeus)
Late Holocene – RS
Butorides striatus – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported this species from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Ardea Linnaeus
477. Ardea sp.
Middle Holocene – SC
Ardea sp. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) associated with this genus 10 bones of at least two individuals from the Congonhas I (SC-LS-30) sambaqui in Tubarão, Santa Catarina.
Threskiornithidae Poche
Theristicus Wagler
478. Theristicus sp.
Quaternary – MG
Ibis ( Theristicus ) sp.; melanopis v. caudata ?; coerulescens ? – Winge 1887: 29–30.
Ibis spec. ( melanopis ) – Goeldi 1894: 559.
Ibis ( Theristicus ) sp. (“ melanopsis var. caudata oder coerulescens ”) – Lambrecht 1933: 735.
Theristicus caudatus – Brodkorb 1963: 279.
Theristicus caudatus [ Ibis ( Theristicus ) sp.] – Mones 1986: 80.
Theristicus caudatus – Cuello 1988: 33.
Theristicus sp. – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported bones of numerous individuals from Lapa da Escrivânia V, represented by the scapula, coracoid, humerus, ulna, radius, carpometacarpus, first phalanx of the major digit of the wing, femur, tibiotarsus, and tarsometatarsus. The bones range from fully formed to fragments of difficult recognition, which he considered to belong to this taxon with a certain degree of uncertainty. Most of them belonged to young birds, which suggests they nested in the area. There is also the coracoid of an adult individual from Lapa da Escrivânia XI.
Winge noted that the bones diverge in many aspects from Plegadis chihi , Platalea leucorodia , and Platalea ajaja . They are very similar to but much larger than Phimosus infuscatus and somewhat smaller than an exceptionally large young specimen of Theristicus melanopis , but otherwise almost completely the same. However, the humerus of T. melanopis is slightly different from the cave material and P. infuscatus .
From its size, Winge considered that the material could belong to Theristicus melanopis , Theristicus caudatus (then still treated as conspecific with the previous taxon), Theristicus caerulescens , or Cercibis oxycerca , and hardly to Mesembrinibis cayennensis . Due to the morphological differences between the fossil material and T. melanopis , T. caerulescens was considered. However, this species, like T. melanopis and C. oxycerca , does not occur (at least nowadays) in the region (Matheu et al. 2017a , 2017b , 2017c ), and, probably for this reason, Brodkorb ( 1963 ) listed the record as T. caudatus .
479. Theristicus caudatus (Boddaert)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material. This is one of the taxa the authors highlighted as no longer living in the region today.
Cathartiformes Seebohm
Cathartidae Lafresnaye
480. Cathartidae indet. 1
Quaternary – MG
Fig. 26 A–C
“un Cathartes plus grand que les espèces actuelles” [?] – Gervais 1844a: 294.
Cathartes [?; in part] – Giebel 1846: 313.
“die Überreste einer hiehergehörigen Art, welche die jetzt dort lebenden an Größe übertraf” [?] – Giebel 1847: 8.
Catharte [sic] [?; in part] – Pictet 1853: 410.
“des Urubus ( Cathartes d’Illiger)” [?] – Liais 1872: 303.
Cathartes aff. papæ [in part] – Lund (in Winge 1887 ): 32.
Gyparchus sp. forma magna; Gyparchus papa et aff. (v. sim.) [“Knogler af en Form, der er meget større end Gyparchus papa og maaske er en egen Art”] – Winge 1887: 14, 32.
“um Urubú-rei muito grande, especie de Sarcoramphus , que excedia muito as dimensões do actual, S. papa ” – Goeldi 1894: 76.
Gypagus papa [in part] – Lambrecht 1933: 744.
Sarcoramphus papa (?) [in part] – Brodkorb 1964: 257.
Sarcoramphus papa [ Gyparchus papa ] [in part] – Mones 1986: 83.
Sarcoramphus papa [in part] – Cuello 1988: 37.
Cathartidae indet. 1 – Nascimento and Silveira 2020: 491.
Along with the description of remains he associated with Sarcoramphus papa , Winge ( 1887 ) described in detail three quite altered fragments apparently of a single individual from Lapa da Escrivânia I, representing a vulture much larger than S. papa but smaller than Vultur gryphus . They are the proximal end of a right humerus that is quite incomplete and partially covered with incrustations (“ Cathartes aff. papæ ” in Lund’s catalog), the proximal end of an ulna, also quite incomplete and partially covered with hardened sediment, and a coracoid in two parts, relatively shorter and more robust than in S. papa . The finding site of the coracoid was not recorded, but it apparently has the same origin of the others based on its aspect and size.
Cathartidae indet. 1: proximal end of the right humerus (A), coracoid in two pieces (B), and proximal end of the ulna (C) (K.L. Hansen); Cathartidae indet. 2: distal half of the left tibiotarsus (MCL A 1795) (D) (C. Cartelle); Pleistovultur nevesi: right tibiotarsus (MHNT-VT-5238), in lateral (E), posterior (F), and medial (G) views (G. Couto Ribeiro). Scale bar: 10 mm.
Winge noted that he was not sure whether these bones belong to an exceptionally large S. papa , a large extinct race, or a vanished species, but that several factors point to the last, and, in this case, it would represent a genus of cathartid other than Sarcoramphus . Alvarenga et al. ( 2008 ) commented that the bones found in Lapa da Escrivânia I were cleaned and re-prepared by Storrs L. Olson and Steven D. Emslie, but they do not show sufficiently preserved structures for a taxonomic diagnosis, and they may belong to the genus Pleistovultur , found in the same region.
These fossils were also mentioned by Lönnberg ( 1902 ) when he described a tarsometatarsus and an incomplete femur of a cathartid from the Pleistocene of the Tarija valley, Bolivia, as Sarcoramphus patruus ( Sarcoramphus here as a synonym of Vultur ). Regarding the possible relationship between the Brazilian and Bolivian remains, he commented (Lönnberg 1902: 8): “Whether this one might be identical with this fossil Condor from Tarija, or not, cannot be decided upon, as unfortunately only remains of bones of the wing of the Brazilian bird are left, as Inspector H. Winge has kindly told me. The size seems, he says, however, to correspond fairly well. But if the Brazilian bird has been a true Gyparchus [= Sarcoramphus ], as I suppose it has, when that judgment has been pronounced by such an able ornithologist as Mr. [Oluf] Winge, it could not, for reasons stated above [differences between the Bolivian fossils and the living king vulture], be identical with the fossil Condor from Tarija”. In the face of the association of a fossil ulna from the Lagoa Santa region to S. papa by Lydekker ( 1891 ; see below), Lönnberg also commented that, although rather scanty, it is right to consider, based on these authorities, the bones from the caves of Brazil as true Sarcoramphus . Agnolín et al. ( 2017a ) assessed the Bolivian material and considered it a valid taxon with an uncertain generic position.
481. † Cathartidae indet. 2
Alvarenga et al. ( 2008 ) reported the distal half of a left tibiotarsus (MCL A 1795) found by Cástor Cartelle in Gruta dos Brejões, Morro do Chapéu, Bahia, the same site where the type specimen of Wingegyps cartellei was found.
Its damaged condyles prevented a better diagnosis of its identity, but the available traits excluded the possibility that it belongs to Vultur , Gymnogyps , Breagyps , Sarcoramphus , Geronogyps , or Pleistovultur , despite the size being close to the last. The authors believed it does not belong to any known genus of Cathartidae, but a new name was not erected in favor of the surfacing of better material.
482. Cathartidae indet. 3
Mayer ( 2013 ) reported cathartid remains from the Lagoa Uri de Cima site in Salgueiro, Pernambuco, found among other vertebrates in a layer dated 13,700–8,000 years BP. It is possible that this material figures among the avian remains reported by Faure and Guérin ( 2013 ).
483. Cathartidae indet. (spp.?) 4
Middle Holocene – RS, SC
Cathartidae indet. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) reported eight indeterminate cathartid bones belonging to at least two individuals from the Congonhas I (SC-LS-30) (Tubarão, Santa Catarina) and Figueira II (Arroio do Sal, Rio Grande do Sul) sambaquis.
Sarcoramphus Duméril
484. Sarcoramphus papa (Linnaeus)
Quaternary – MG
Fig. 27 A–H
Cathartes aff. papæ [in part] – Lund (in Winge 1887 ): 32.
Gyparchus papa , typicus; Gyparchus papa et aff. (v. sim.) [“Knogler, der passe til Gyparchus papa ”] – Winge 1887: 14, 32–33.
Vulturides [in part] – Goeldi 1894: 76.
Gypagus papa [in part] – Lambrecht 1933: 744.
Sarcoramphus papa [in part] – Brodkorb 1964: 257.
Sarcoramphus papa [ Gyparchus papa ] [in part] – Mones 1986: 83.
Sarcoramphus papa [in part] – Cuello 1988: 37.
Sarcoramphus papa [in part] – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported remains of two individuals from Lapa do Baú. One of them (whose most elements were identified as “ Cathartes aff. papæ ” in Lund’s catalog) is represented by the maxilla, jugal arches, tip of the mandible, part of the anterior portion of the sternum, scapula, humerus, ulna, radius, and carpometacarpus. They closely match recent bones of this species, presenting only small variations, of which the most notable is the more robust ulna. Of the other individual, there are only the fragmented remains of the proximal end of the left humerus and the distal half of the right tibiotarsus, slightly larger than in recent bones, where the humerus also presents some variations, especially in the much thicker head.
Sarcoramphus papa , unspecified Lagoa Santa locality: proximal end of the left ulna (NHMUK PV OR 18887), dorsal (A), caudal (B), cranial (C), ventral (D), proximal (E), and distal (F) views (S. Chapman); Lapa do Baú: fragment of the proximal end of the left humerus (G) and distal half of the right tibiotarsus (H). Wingegyps cartellei , Lapa do Tiú, paratypes: proximal half the right ulna (ZMUC 1118), dorsal (I) and ventral (J) views; distal end of the right humerus (ZMUC 1116), caudal (K) and cranial (L) views (K.L. Hansen); Morro do Chapéu: incomplete neurocranium (MCL A 782, holotype), dorsal (M) and ventral (N) views; distal end of the left humerus (MCL A 1678) (O); left humerus (P) (MCL A 670) (C. Cartelle). Scale bar: 10 mm.
Lydekker ( 1891 ) reported the proximal end of a left ulna (NHMUK PV OR 18887), indistinguishable from those of recent skeletons. It came from the Lagoa Santa region and was acquired by the British Museum (Natural History) in 1848 as part of the Claussen Collection.
Vultur Linnaeus
485. Vultur gryphus Linnaeus
Alvarenga ( 1996 , 1998 ) associated a premaxilla (MNRJ-A-LV-82) discovered by the Franco-Brazilian Archeological Mission (1971–1976) in Lapa Vermelha IV in Confins, Minas Gerais, with Vultur gryphus , a species never positively recorded in Brazil (Pacheco et al. 2021 ). Previously, he alluded to it briefly in Sick ( 1993 , 1997 ).
Alvarenga pointed out that the presence of this taxon at the beginning of the Holocene in Minas Gerais might indicate that the climate of the region was colder, with strong wind currents that would facilitate its takeoff and flight maintenance, or even that its distribution during this time spanned over a broad area of the continent’s east coast. The sediments where the bone was found are 10,220–11,680 years old.
Agnolín ( 2016b ), Agnolín et al. ( 2016 ), and Agnolín et al. ( 2017a ), reviewing the fossil records associated with V. gryphus , restricted their occurrence to the late Pleistocene of northwestern Peru and southern Chile. They noted that, although the Brazilian specimen is similar to this species, its fragmentary nature and the lack of knowledge of the beak morphology in most South American fossil condors prevent its determination, and it is not unlikely that it may belong to Pleistovultur nevesi .
Coragyps Le Maout
486. Coragyps atratus (Bechstein)
Quaternary – MG, RN
Catharistes atratus – Winge 1887: 31, 8.
Vulturides [in part] – Goeldi 1894: 76.
Catharistes urubu – Lambrecht 1933: 744.
Coragyps atratus – Brodkorb 1964: 257.
Coragyps atratus [ Catharistes atratus ] – Mones 1986: 82–83.
Coragyps atratus – Cuello 1988: 36–37.
Coragyps atratus – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported bones of several individuals, young and adults, from Lapa da Escrivânia V, and those of apparently a single individual from Lapa da Lagoa do Sumidouro. The material is represented by the quadrate, coracoid, humerus, ulna, carpometacarpus, the first phalanx of the major digit of the wing, femur, tibiotarsus, tarsometatarsus, and the first phalanx of digit III of the foot. Winge noted that Coragyps atratus and Cathartes aura could be easily distinguished in most bones.
Coragyps atratus – Costa et al. 2023b: 35.
Costa et al. ( 2023b ) reported a fragment of coracoid among material found in two sites (Ravina do Leon and Ravina das Araras) of the Lajedo de Soledade karstic area in Apodi, Rio Grande do Norte.
Cathartes Illiger
487. Cathartes aura (Linnaeus)
Winge ( 1887 ) reported some possibly associated leg bones from Lapa da Escrivânia V, bones of apparently a single individual from Lapa da Escrivânia XI, and two tarsometatarsi from Lapa da Lagoa do Sumidouro, from an adult and a young specimen. The total material is represented by the scapula, coracoid, humerus, ulna, carpal [27] , tibiotarsus, and tarsometatarsus.
Kneip et al. ( 1989b , 1994 ) reported two bones of at least one individual from layer II (4,160±180 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Costa et al. ( 2023a ) reported the proximal end of a right carpometacarpus of quaternary age from Ravina das Araras in the karstic area of Lajedo de Soledade in Apodi, Rio Grande do Norte. The fossil was collected in 2019 by a joint fieldwork of Laboratório de Paleontologia of Faculdade de Geologia of UERJ and researchers of UERN and UFRJ. Later, Costa et al. ( 2023b ) further associated with this species a tibiotarsus fragment from the same area, without specifying if it is from Ravina das Araras or Ravina do Leon.
Cathartes aura – Leoni et al. 2024: 3, figs 2–3.
Leoni et al. ( 2024 ) reported several elements from Toca da Boa Vista in Campo Formoso, Bahia. The material comprises a cranial fragment (LEG 2288), three cervical vertebrae (LEG 2890, LEG 2891, and LEG 2892), sternum (LEG 2297), furcula (LEG 2295), distal fragment of the left coracoid (LEG 2303), distal fragment of the right coracoid (LEG 2294), scapula (LEG 2305), distal fragment of the left humerus (LEG 2301), proximal fragment of the right humerus (LEG 2296), distal fragment of the left ulna (LEG 2300), distal fragments of the right ulna (LEG 2308 and LEG 2312), a well-preserved right carpometacarpus (LEG 2311), synsacrum (LEG 2289), proximal fragment of the left femur (LEG 2307), fragments of the right tibiotarsus (LEG 2310 and LEG 2313), and proximal fragment of the right tarsometatarsus (LEG 2293). The bones were found disarticulated in a corridor about 2 to 2.5 kilometers from the cave entrance, where the individual probably died. Markings present in the LEG 2308 ulna and the LEG 2310 tibiotarsus were associated with the ichnospecies Nihilichnus nihilicus and were likely made by a small felid such as Leopardus tigrinus or Herpailurus yagouaroundi .
†Brasilogyps Alvarenga
488. † Brasilogyps faustoi Alvarenga
Late Oligocene/Early Miocene – SP
Fig. 32 C
Type locality: Taubaté sedimentary basin: municipality of Tremembé, state of São Paulo, Brazil.
Etymology: Brasilogyps , with the Greek gyps , “Brazilian vulture”, and faustoi , honoring Brazilian paleontologist Fausto Luiz de Souza Cunha (1926–2000) (MN), for his work done in the Taubaté Basin and support given to Alvarenga’s research.
Brasilogyps faustoi , sp. n. [new genus and species] – Alvarenga 1985b: 349–357, figs 1–2, 3A, 4, 5A, 6C, 7A.
Brasilogyps faustoi – Mones 1986: 82.
Brasilogyps faustoi – Cuello 1988: 11.
Brasilogyps faustoi – Alvarenga 1993a: 62.
Brasilogyps faustoi – Alvarenga 1997: 123.
Brasilogyps faustoi – Kellner 1998: 654, 659.
Brasilogyps faustoi – Alvarenga and Höfling 2000: 589.
Brasilogyps faustoi – Alvarenga and Höfling 2004: 827.
Brasilogyps faustoi – Alvarenga and Höfling 2011: 127.
Brasilogyps faustoi – Carmo et al. 2024: 5, 3C–D.
Alvarenga ( 1985b ) described this new genus and species from two associated bone fragments collected in January 1978 in the montmorillonite clays of the Tremembé Formation in the Fazenda Santa Fé.
The material (MN 4045-V, holotype) consists of the distal end of a right tibiotarsus with a significantly worn extremity and the proximal end of a right tarsometatarsus. The study of the material demonstrated a greater proximity to Coragyps , being slightly larger and more robust than Coragyps occidentalis , from the Quaternary of North America, which was larger than the living Coragyps atratus (Alvarenga 1985b , Brodkorb 1964 ).
Its age and geographical origin are valuable for understanding the evolution of cathartids. It is the oldest known cathartid from South America (Alvarenga 1993b , 1997 , Alvarenga et al. 2008 , Tambussi and Degrange 2013 , Mayr 2016 ), or even in the Americas as a whole, if Phasmagyps patritus from the Eocene of Colorado is not in fact a member of this family, as noted by Olson ( 1985b ) regarding its problematic systematic position.
†Wingegyps Alvarenga & Olson
489. † Wingegyps cartellei Alvarenga & Olson
Quaternary – BA, MG
Figs 27I–P, 35D
Type locality: Brazil, Bahia State, Município de Morro do Chapéu, Gruta dos Brejões (11°00’30”S, 41°26’07”W), elevation ca. 600m.
Etymology: Wingegyps , with the Greek gyps , “Winge’s vulture”, honoring ornithologist Oluf Winge for his perspicacity in recognizing the distinctiveness of the fossils, and cartellei , honoring Spanish-Brazilian paleontologist Cástor Cartelle Guerra (PUC Minas), in recognition for his excavations at Gruta dos Brejões and his contributions to Brazilian paleontology.
Cathartes – Lund (in Winge 1887 ): 33.
G. sp. indet. magnitudine Catharistæ atrati – Winge 1887: 33–34, 7.
“um Urubú do tamanho de Cathartes aura ” [?] – Goeldi 1894: 76.
“einen angeblich ausgestorbenen Vertreter der Cathartidae” – Lambrecht 1921: 29.
Cathartidarum gen. sp. indet. [in part] – Lambrecht 1933: 397.
Gen. sp. indet. – Lambrecht 1933: 744.
Catharistes urubu (?) – Lambrecht 1933: 744.
Cathartidarum – Fisher 1944: 294.
Coragyps atratus (?) – Brodkorb 1964: 257.
Wingegyps cartellei , new species [new genus and species] [in part] – Alvarenga and Olson 2004: 1–9, figs 3–4.
Wingegyps cartellei [in part] – Alvarenga and Höfling 2011: 127.
“ Wingegyps (Amazonian Vulture)” – Campbell 2016: 120–121.
Wingegyps cartellei – Nascimento and Silveira 2020: 491, 5A–C.
Winge ( 1887 ) described in detail two bones of similar aspect (determined as “ Cathartes ” in Lund’s catalog) from Lapa do Tiú [35] , the distal half of a right humerus (ZMUC 1116) and the proximal end of a right ulna (ZMUC 1118). The humerus notably diverges from the known living cathartids, but Winge did not attribute the material to a new genus or species and referred to it just as a cathartid of similar size to Coragyps atratus . An illustration of the humerus was included, in which it was compared with a fossil humerus of C. atratus also found by Lund.
Before the description by Alvarenga and Olson (see below), there was some confusion regarding how this material was referred to. Lambrecht ( 1933: 397) mentioned the humerus as “Cathartidarum gen. sp. indet.” (“undetermined cathartid genus and species”) and even listed unrelated material from France with a similar provisional term (“Cathartidarum gen. inc.”). The same material from Lapa do Tiú was mentioned as “Gen. sp. indet.” of Cathartidae in the same study (Lambrecht 1933: 744). The first term, however, was used as a genus by Fisher ( 1944: 294), who wrote “ Cathartidarum is the genus erected for a Pleistocene humerus from Lagoa Santa, Brazil, by Winge (1888)” and “It is about the size of ’ Cathartes atratus’ ”. Although he did not cite Lambrecht’s study, the mentioned information is the same found there. He also noted (Fisher 1944: 295) that, if it is a valid genus, “it is the oldest member of the King Vulture group, and is probably a close relative of S. papa ”. Fisher based his assumption of affinity on Miller ( 1931 ), who only noted that one species close to the king vulture was found in Brazilian Pleistocene cave deposits. Miller did not specify which cathartid remains described by Winge he was referring to, but it was likely the fossils determined as Gyparchus papa et aff. (v. sim.) and not the material mentioned by Lambrecht. Tordoff ( 1959 ) kept the name used by Fisher and the same interpretation of Miller’s text, although he noted he did not consult Winge’s study. The name was also kept in publications (Wing 1956 , Stager 1964 ) that reused the phylogenetic tree presented by Fisher. Despite predating Wingegyps , the circumstances in which Fisher used the name as a genus for the humerus from Lapa do Tiú are not valid according to the norms of the International Commission on Zoological Nomenclature ( 2000: Art. 13.3). Emslie ( 1988 ) mentioned “ Cathartidarum ” as a “species” that is no longer considered a member of this family when commenting on the use of fossil species in systematics analysis of Cathartidae (including Fisher’s).
Alvarenga and Olson ( 2004 ), while identifying bird bones from Gruta dos Brejões in Morro do Chapéu, Bahia, noticed an ovoid-shaped skull similar to that of Gymnogyps , but much smaller, and two humeri, probably belonging to the same species as the skull, of which a well-preserved distal fragment appeared to be identical to the specimen illustrated by Winge ( 1887 ). The reanalysis of the bones discovered by Lund confirmed that this is indeed a new taxon, and, along with the inclusion of the material from Bahia, the genus and species Wingegyps cartellei were erected.
The material found in Bahia comprises an incomplete neurocranium (MCL A 782, holotype), a complete but very worn left humerus (MCL A 670), and the distal third of another left humerus (MCL A 1678). It was dated about 12,200 years BP based on that of a giant sloth coprolite found in the same locality and associated mammalian megafauna found in the caves of Bahia and Minas Gerais.
The species is closer to condors, especially Gymnogyps , than to the smaller genera of the family (i.e., Cathartes and Coragyps ), which are basically differentiated by skull morphology (Alvarenga and Olson 2004 ). However, it is much smaller than any known condor and slightly smaller than Cathartes burrovianus , the smallest living cathartid. The length from the tip of the beak to the end of the tail was estimated at 50 cm and the wingspan at 130 cm (Pivetta 2003 ). The discovery of Wingegyps highlights that the condors had a much greater range of sizes in the past.
Its distribution range was probably much wider than the extent of the two geographical points where its remains are known. Had it been collected in any other deposit, it might have been treated as Cathartes or Coragyps due to its reduced size. Three complete coracoids, two incomplete humeri, and one complete humerus of an undescribed Wingegyps species were reported by Steadman et al. ( 2015 ) from the late Pleistocene of the asphalt deposit of Mene de Inciarte in Mara, Zulia, northwestern Venezuela.
Alvarenga and Olson postulated that W. cartellei could hardly process most of the carcasses or compete for them with other vulture species because of its small size. If, like other condors, it did not possess the olfactory capabilities of the Cathartes species, it would also have had difficulty competing with them for smaller carcasses. It could, however, have harvested the fruits of palm trees, a niche best exploited in the Old World by the accipitrid Gypohierax angolensis in Africa, which consumes the soft mesocarp of the African oil palm Elaeis guineensis . This niche is also present in Brazil through the consumption of the fruits of this same species (which was introduced) by Cathartes aura , which also feeds on the macaúba palm Acrocomia aculeata , a native species whose broad geographical distribution coincides with the known past range of W. cartellei . The authors added that, although this small condor may have occupied this niche, its habits may have been similar to those of the accipitrid Neophron percnopterus , which consumes scraps of carcasses left by larger vultures. This behavior could explain its extinction since many New World scavenging birds disappeared alongside the mammalian megafauna.
†Pleistovultur Alvarenga, Brito, Migotto, Hubbe & Höfling
490. †Pleistovultur nevesi Alvarenga, Brito, Migotto, Hubbe & Höfling
Quaternary – MG, RN
Figs 26E–G, 35C
Type locality: Brazil, Minas Gerais State, Municipality of Matozinhos, Gruta Cuvieri (19º28’36”S, 44º00’41”W), elevation ca. 812m.
Etymology: Pleistovultur , a portmanteau of Pleistocene and the genus Vultur , “Pleistocene condor”, and nevesi , honoring Brazilian anthropologist Walter Alves Neves (LEEH-IB-USP), responsible for the team that collected and forwarded the fossil to the authors.
“new extinct genus” – Migotto and Alvarenga 2007: 29R.
Pleistovultur nevesi sp. nov. [new genus and species] – Alvarenga et al. 2008: 613–618, 2.
Pleistovultur nevesi – Alvarenga and Höfling 2011: 127.
Pleistovultur nevesi – Nascimento and Silveira 2020: 491, 5D–E.
Alvarenga et al. ( 2008 ) described this new genus and species of large condor from a right tibiotarsus (MHNT-VT-5238) collected by Alex Hubbe around July 2005 in the Locus 1 of Gruta Cuvieri (A. Hubbe, personal communication) in Matozinhos, Minas Gerais. The bone is well-preserved and apparently belongs to an adult individual, 25% larger than in Sarcoramphus and 11% smaller than in Vultur . Its existence was first alluded to by Migotto and Alvarenga ( 2007 ).
The assignment of this material to the late Pleistocene or the early Holocene was defined based on associated faunal remains present in the same cave (two Scelidodon ground sloths were dated about 9,990 and 12,510 years BP) and adjacent ones previously dated since the taphonomic conditions of the specimen could not be adequately determined due to previous work carried out on the site. Other materials possibly referable to this taxon were described in the literature (see Cathartidae indet. 1 and Vultur gryphus ).
Its discovery adds to the great diversity of cathartids in South America during the Pleistocene, and the disappearance of the megafauna has certainly led many of these species to extinction.
Costa et al. ( 2023a ) associated with this species the distal end of a right tibiotarsus of quaternary age from Ravina do Leon in the karstic area of Lajedo de Soledade ( Fig. 1 .18) in Apodi, Rio Grande do Norte. The fossil was collected in 1993 by Prof. Leon Diniz Dantas de Oliveira and his staff and considerably expanded the species’ known distribution range.
† Teratornithidae Miller
†Taubatornis Olson & Alvarenga
491. † Taubatornis campbelli Olson & Alvarenga
Late Oligocene/Early Miocene – SP
Figs 28, 32D
Type locality: Brazil, São Paulo State, 2 km NE of Tremembé, Santa Fé Farm (22°30’S, 45°32’W).
Etymology: Taubatornis , with Greek ornis , “bird of Taubaté”, and campbelli , honoring American paleontologist Kenneth E. Campbell, Jr. (1943–) (LACM), in recognition of his contributions to the knowledge of the teratornithids.
Taubatornis campbelli , new species [new genus and species] – Olson and Alvarenga 2002: 701–705, figs 1–2.
Taubatornis campbelli – Alvarenga and Höfling 2011: 127.
Taubatornis campbelli – Carmo et al. 2024: 5, 3E–F.
Olson and Alvarenga ( 2002 ) described this new genus and species from two fragments from the montmorillonite clays of the Tremembé Formation of the Fazenda Santa Fé, about 4 m below the uppermost level of the shales. It is the smallest teratornithid known so far, but a large bird nevertheless, and the only one described for the Brazilian territory. Its wingspan was estimated at 1.9 m (Pivetta 2003 ).
The material consists of the distal end of a right tibiotarsus (MHNT-VT-5154, holotype) and the proximal end of a left ulna lacking part of the olecranon (MHNT-VT-5155, paratype).
As observed with other teratornithid occurrences, it was found syntopic with cathartid fossils ( Brasilogyps faustoi ). The shales of the lacustrine environment of the Tremembé Formation suggest an alternation between wet and dry seasons, resulting in the periodic mortality of large numbers of small fishes, which may have been responsible for the presence of these birds.
It constitutes the oldest record for the family, about 25 million years older than the next oldest member, the 6 million years old gigantic Argentavis magnificens of the Miocene of Argentina, and reinforces the theory that this family originated in South America (Olson and Alvarenga 2002 , Agnolín 2016b ).
Accipitriformes Bonaparte
492. Accipitriformes indet.
Quaternary – RN
Accipitriformes indet. – Costa et al. 2023b: 35.
Costa et al. ( 2023b ) associated with an accipitriform two ungual phalanges from material found in two sites (Ravina do Leon and Ravina das Araras) of the Lajedo de Soledade karstic area in Apodi, Rio Grande do Norte.
Taubatornis campbelli: distal end of the right tibiotarsus (MHNT-VT-5154, holotype), anterior (A) and posterior (B) views; proximal end of the left ulna (MHNT-VT-5155, paratype), dorsal (C) and ventral (D) views (G. Couto Ribeiro). Scale bar: 10 mm.
Accipitridae Vigors
493. Accipitridae indet. 1
Quaternary – MG
Thrasaëtus ? sp. – Winge 1887: 35–36.
Buteonines [in part] – Goeldi 1894: 76.
Thrasaëtus sp. – Lambrecht 1933: 747.
Harpia harpyja – Brodkorb 1964: 283.
Harpia harpyja [ Thrasaetus ? sp.]- Mones 1986: 84.
Harpia harpyja – Cuello 1988: 39.
Harpia harpyja – Penido et al. 2012: 8.
Harpia harpyja – Penido et al. 2013: 117.
Accipitridae indet. 1 – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported the distal end of an ulna of unknown origin. He noted it is somewhat larger than in Haliaeetus albicilla , Aquila chrysaetos , and Trigonoceps occipitalis , smaller than in Gyps fulvus , and close to Gypaetus barbatus . Its shape is very close to that of Buteo and even more so to that of Spizaetus ( S. ornatus and S. tyrannus ). He concluded that, of the living South American species, one can think of Harpia harpyja and Urubitinga coronata , although only the first was known from the Lagoa Santa region.
494. Accipitridae indet. 2
Quaternary – MG
Buteonin. indet., s. Tillæg til Buteoninæ [“Løb (nederste Ende) og Mellemhaand (øverste Ende); en god Del større end hos Buteo ( Asturina ) nattereri ”] – Winge 1887: 15, 36.
Buteo sp. [in part] – Lambrecht 1933: 748.
Accipiter sp. [?; in part?] – Mones 1986: 83.
Accipitridae indet. 2 – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported the distal end of a tarsometatarsus and the proximal end of a carpometacarpus from “various caves” [26] . He noted that they are significantly larger than those of Rupornis magnirostris nattereri .
495. Accipitridae indet. 3
Quaternary – MG
Buteonin. sp. indet., s. Anm. til Buteoninæ [“Et Albueben (Stump) og et Taaled af en Art af lignende Størrelse som foregaaende”] – Winge 1887: 13, 36.
Buteo sp. [in part] – Lambrecht 1933: 748.
Accipiter sp. [?; in part?] – Mones 1986: 83.
Accipitridae indet. 3 – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported the fragment of an ulna and a pedal phalanx from Lapa da Lagoa do Sumidouro, similar in size to the previous one.
496. Accipitridae indet. 4
Winge ( 1887 ) reported a pelvis and a coracoid from Lapa da Escrivânia V, roughly equivalent in size to the previous two. Besides these three indeterminate accipitrids he described along with the other remains of this family, he mentioned the existence of several other fragments.
497. Accipitridae indet. (spp.?) 5
Carvalho ( 1984 ) reported three accipitrid ulnae, worked in two different manners, among the material discovered in 1978 at the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
498. Accipitridae indet. 6
Queiroz and Chaix ( 1999 ) reported remains of a large bird of prey, “possibly a harpy eagle”, found in 1998 in association with adult male human remains (Carvalho et al. 2002 ) in a large ceramic pot from burial 34 of the Justino site in Canindé de São Francisco, Sergipe. They described the material as a dissociated cranium and beak, a humerus and a tarsometatarsus found over the abdomen of the human body, and another humerus buried vertically in the sediment. Queiroz and Carvalho ( 2005 ) reported it to consist of cranial bones, the beak, the left humerus, the left femur, and the left tarsometatarsus. The material was radiocarbon-dated at 1770±60 years and is deposited in the MAX collection. By the size of the bones, this specimen and the one described below are referred to as accipitrids here despite being called “falconids”.
499. Accipitridae indet. 7
Carvalho et al. ( 2002 ) briefly mentioned and depicted articulated remains of a medium-sized bird of prey found in burial 166 of the Justino site in Canindé de São Francisco, Sergipe. It was found in 1998, in the body of an adult human of indeterminate sex. The beak, a coracoid, the wing bones, some ribs, the pelvic girdle, and some hindlimb bones were later identified (Queiroz and Carvalho 2005 ). Cardoso ( 2015 ) noted that a deeper analysis of the material was not possible as it would compromise its integrity. It was radiocarbon-dated at 1770±60 years (Queiroz and Carvalho 2005 ) and is deposited in the MAX collection.
Besides these accipitrid and ciconiid remains (see Ciconiidae indet. 3), two humeri and a tibiotarsus of another bird were found in burial 131, but they could not be determined due to the lack of comparative material (they were depicted in figure 34 of Queiroz and Chaix 1999 ). Additionally, remains of a mustelid were found in similar association with human remains in burial 119. The presence of these animals in the burials may indicate a ritual connotation, possibly as psychopomps, and constitute the only known cases of this nature in Brazil (Queiroz et al. 2017 ).
500. Accipitridae indet. 8
Quaternary – BA
Accipitridae – Silva and Cozzuol 2010: 111.
Silva and Cozzuol ( 2010 ) associated with this family an incomplete tarsometatarsus lacking the proximal epiphysis among material found in the 1980s at Toca da Boa Vista in Campo Formoso, Bahia. The fossil is deposited in the MHNJB/UFMG collection.
501. Accipitridae indet. 9
Quaternary – CE
Fig. 18 D–F
“Neornithes: indeterminadas” [in part] – Souza Cunha 1961: 5.
“Neornithes, Ordens e gêneros indeterminados” [in part] – Paula Couto 1961: 8.
“aves (indeterminadas)” [?; in part] – Paula Couto 1962: XIX.
“Aves Neornithes” [?; in part] – Paula Couto 1978b: 429.
“Neognathae, Undeterminated” [?; in part] – Paula Couto 1980: 145.
“Aves neornites” [?; in part] – Silva Santos 1982: 118.
Neognathae indet. [?; in part] – Ximenes 2009: 472.
Neognathae indet. [?; in part] – Souto and Carvalho 2010: 116.
“gavião fóssil” [in part] – Metello and Araújo Júnior 2012: 18.
“gavião fóssil” [in part] – Metello and Araújo Júnior 2013: 69–70.
Acciptriformes indet. [sic] [in part] – Araújo Júnior 2015: 193.
Acciptriformes indet. [sic] [in part] – Araújo Júnior 2016: 150, 2B.
“bird remains” [in part] – Patusco et al. 2016a: 190.
“aves” [in part] – Patusco et al. 2016b: 274.
Acciptriformes indet. [sic] [in part] – Waldherr et al. 2017: 471.
Acciptriformes indet. [sic] [in part] – Waldherr et al. 2019: 115.
Accipitridae indet. – Costa et al. 2024: 2, 2A, D–E, H–I.
Metello and Araujó Junior ( 2012 , 2013 ) associated with a hawk remains of pleistocenic age found in tank 2 of the João Cativo locality in Itapipoca, Ceará. They initially attributed to a single individual four well-preserved cervical vertebrae (MN 3326-V [36] ), the distal end of a right tarsometatarsus (MN 3275-V), a complete ungual phalanx (MN 3263-V), a hallux (MN 3265-V [37] ), and still-indeterminate fragments (MN 3270-V, MN 3271-V, MN 3272-V, MN 3273-V, MN 3293-V, and MN 3294-V).
The mentioned material represents multiple, disparate species, and part of it does not belong to birds. Costa et al. ( 2024 ) attributed to Accipitridae only the ungual phalanx MN 3263-V and phalanges MN 3266-V, MN 3267-V, and MN 3268-V; the fragment of tarsometatarsus MN 3275-V was assigned to Rhea americana .
502. Accipitridae indet. (spp.?) 10
Holocene – PR
Accipitridae indet. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) reported two indeterminate accipitrid bones from the Toral 51 and Guaraguaçu B sambaquis of Paranaguá and Pontal do Paraná, respectively, in Paraná.
Gypaetinae Bonaparte
Chondrohierax Lesson
503. Chondrohierax uncinatus (Temminck)
Quaternary – MG
Cymindis uncinatus – Winge 1887: 35.
Buteonines [in part] – Goeldi 1894: 76.
Cymindis ( Regerhinus ) uncinnatus – Lambrecht 1933: 751.
Chondrohierax uncinatus – Brodkorb 1964: 288.
Chondrohierax uncinatus – Mones 1986: 84.
Chondrohierax uncinatus – Cuello 1988: 37.
Cymindis uncinatus – Penido et al. 2012: 8.
Cymindis uncinatus – Penido et al. 2013: 117.
Chondrohierax uncinatus – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported a right femur and a left tibiotarsus of unknown origin. The tibiotarsus corresponds well to Chondrohierax uncinatus , although insignificantly larger and more robust and presenting small, probably individual variation. Leptodon cayanensis has a similar but larger and slightly more robust tibiotarsus. The femur, of which there was no comparative correspondent of C. uncinatus , is very similar to that of L. cayanensis , except for the much smaller size. Winge also noted that the tibiotarsus of Elanoides forficatus is different but still close, and Elanus and Ictinia are very different from the preceding.
Accipitrinae Vigors
Accipiter Brisson
504. Accipiter sp.
Quaternary – MG
Accipiter sp., magnitudine A. nisi ♀ , forte A. pileatus ♂ – Winge 1887: 36.
Buteonines [in part] – Goeldi 1894: 76.
Accipiter sp. – Lambrecht 1933: 749.
Accipiter sp. [in part?] – Mones 1986: 83.
Accipiter bicolor – Penido et al. 2012: 8.
Accipiter bicolor – Penido et al. 2013: 117.
Accipiter sp. – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) determined the proximal and distal ends of a right tarsometatarsus of unknown origin as belonging to a bird of this genus, of size comparable to that of a female Accipiter nisus , probably a male Accipiter bicolor pileatus .
505. Accipiter bicolor (Vieillot)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Geranospiza Kaup
506. Geranospiza caerulescens (Vieillot)
Penido et al. ( 2012 , 2013 ) reported a right tarsometatarsus, a fragmented left tarsometatarsus, and a synsacrum among material collected in 1980 and 1984 by the staff of the Laboratório de Paleontologia of MCL PUC Minas at Gruta dos Brejões, in Morro do Chapéu, Bahia. The material was associated with this species by the MHNT staff.
Heterospizias Sharpe
507. Heterospizias meridionalis (Latham)
Holocene – RJ
Heterospizias meridionalis – Kneip et al. 1994: 50.
Kneip et al. ( 1994 ) reported three bones of at least one individual from layer I (3,800±190 years BP), two bones of at least one individual from layer II (4,160±180 years BP), and five bones of at least one individual from layer IV (4,520±190 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Urubitinga Lafresnaye
508. Urubitinga urubitinga (Gmelin)
Holocene – RJ
Buteogallus urubitinga – Kneip et al. 1994: 50.
Kneip et al. ( 1994 ) reported a bone from layer II (4,160±180 years BP) and another from layer IV (4,520±190 years BP) of Sambaqui da Beirada in Saquarema, Rio de Janeiro.
Rupornis Kaup
509. Rupornis magnirostris (Gmelin)
Quaternary – MG, SC
Buteo ( Asturina ) nattereri – Winge 1887: 36.
Buteonines [in part] – Goeldi 1894: 76.
Buteo ( Asturina ) Nattereri – Lambrecht 1933: 748.
Buteo magnirostris – Brodkorb 1964: 283.
Buteo magnirostris [ Buteo ( Asturina ) nattereri ] – Mones 1986: 84.
Buteo magnirostris – Cuello 1988: 38.
Buteo natteri [sic] – Penido et al. 2012: 8.
Buteo natteri [sic] – Penido et al. 2013: 117.
Rupornis magnirostris nattereri – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported a humerus and a tibiotarsus from Lapa da Escrivânia V, a coracoid that almost certainly belongs to this species from Lapa da Escrivânia XI, a radius from Lapa da Lagoa do Sumidouro, and a tibiotarsus of a very young individual of recent age [28] .
Rupornis magnirostris – Gilson and Lessa 2021: 226.
Gilson and Lessa ( 2021 ) reported two bones belonging to at least one individual from the second occupation and another element from the third occupation of the Rio do Meio site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
Geranoaetus Kaup
510. Geranoaetus albicaudatus (Vieillot)
Holocene – RJ
Buteo albicaudatus – Kneip et al. 1994: 50.
Kneip et al. ( 1994 ) reported this species among the material from sambaquis of Beirada, Moa, and Pontinha in Saquarema, Rio de Janeiro, without specifying the exact site or the number of remains.
511. Geranoaetus melanoleucus (Vieillot)
Quaternary – MG
Buteo sp., melanoleucus v. aff., s. under B. melanoleucus [in part] – Winge 1887: 14.
Buteo melanoleucus – Winge 1887: 36.
Buteonines [in part] – Goeldi 1894: 76.
Buteo melanoleucus – Lambrecht 1933: 748.
Buteo fuscescens – Brodkorb 1964: 281.
Buteo fuscescens [ Buteo melanoleucus ] – Mones 1986: 83–84.
Geranoaetus melanoleucus – Cuello 1988: 38.
Geranoaetus melanoleucus – Penido et al. 2012: 8.
Geranoaetus melanoleucus – Penido et al. 2013: 117.
Geranoaetus melanoleucus – Nascimento and Silveira 2020: 491.
Winge ( 1887 ) reported several bones of an individual (and some of a second) from Lapa da Escrivânia V. The material consists of the neurocranium, mandibular branch, thoracic vertebra, part of the synsacrum and pelvis, scapula, humerus, ulna, radius, carpal [27] , carpometacarpus, femur, tibiotarsus, first phalanx of the hallux, second phalanx of digit II, and ungual phalanx. They correspond precisely to the skeleton of a Chilean specimen of Geranoaetus melanoleucus and are larger and divergent than Geranoaetus albicaudatus . There are femora and tibiotarsi of two individuals, one of them slightly larger than in a recent skeleton. There is also a third phalanx of digit III from the same site, somewhat small for this species. The proximal end of a humerus from Lapa da Escrivânia XI fits well into this species. A radius and several pedal phalanges (including the second of the right and left digits II) from Lapa da Escrivânia III possibly belong to this or a close species. They are slightly smaller than in G. melanoleucus and somewhat larger than and less similar to G. albicaudatus . Winge also noted that Spizaetus ornatus and Spizaetus tyrannus , of comparable size, differ in several ways.
Buteo Lacépède
512. Buteo sp.
Rosa ( 2004 ) reported two bones of different individuals from the Paranaíba phase square 12H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
Strigiformes Wagler
513. Strigiformes indet.
Silva ( 2013 , 2014 ) reported strigiform material among remains collected by the staff of NEA-UFPE, starting in the 1990s, in rock shelters of the Pedra do Alexandre site in Carnaúba dos Dantas, Rio Grande do Norte.
Tytonidae Mathews
Tyto Billberg
514. Tyto furcata (Temminck)
Quaternary – MG, PI, RS
Strix perlata [in part] – Lund 1837: 247–248.
Strix [?; in part] – Gervais 1844a: 294.
Strix [?; in part] – Giebel 1846: 313.
Strix [?; in part] – Giebel 1847: 11.
Strix [?; in part] – Pictet 1853: 411.
“les Strix” [?; in part] – Liais 1872: 303.
Strix flammea ( perlata ) [in part?] – Winge 1887: 37.
Strix flammea ( perlata ) – Goeldi 1894: 76.
Strix flammea ( perlata ) [in part?] – Lambrecht 1933: 770.
Tyto alba [in part?] – Brodkorb 1971: 232.
Tyto alba [ Strix flammea ] [in part?] – Mones 1986: 96.
Tyto alba [in part?] – Cuello 1988: 55.
Tyto furcata [in part?] – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported bones in large numbers from many individuals (ranging from very young to adults) from Lapa da Escrivânia V, two bones from Lapa da Escrivânia XI, a bone from Lapa da Lagoa do Sumidouro, several from an unknown origin, and some more of recent age [28] . The material is represented by various skull parts, scapula, coracoid, humerus, ulna, radius, carpometacarpus, first phalanx of the major digit of the wing, pelvis, femur, tibiotarsus, fibula, tarsometatarsus, and several pedal phalanges, with noticeable individual variation.
Lund ( 1837 ) reported bones of varying ages and numerous traces of its predatory activities (on rodents and bats) in several parts of Lapa Nova de Maquiné. Perhaps they figure among the material of unknown origin and of recent age mentioned by Winge. Lund ( 1841a ) also associated with them the mounds of remains of small mammals and birds, of varying ages, from the very first cave he studied, near the village of Cachoeira do Campo. He mentioned remains of the owl’s feeding activities in the caves several times in further works (Lund 1839 , 1841b , 1841c , 1846 ).
Guérin et al. ( 1993a , 1993b , 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least three adults and one young individual are present in the material.
Jacobus ( 2004 ) reported remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
Strigidae Leach
515. Strigidae indet. 1
Quaternary – MG
Syrnium sp. e minoribus – Winge 1887: 38.
“duas especies de Syrnium , das quaes uma maior, outra menor” [in part] – Goeldi 1894: 76.
Syrnium sp. (“Mehrere Formen”) [in part] – Lambrecht 1933: 771.
Syrnium sp. [in part] – Mones 1986: 96.
Strigidae indet. 1 – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported a tarsometatarsus lacking the proximal end from Lapa da Escrivânia XI. He noted it matches very closely the tarsometatarsus of an indeterminate “ Syrnium ” from the Lagoa Santa region, which may be “ Syrnium melanotum ” (= Pulsatrix melanota or Pulsatrix koeniswaldiana ? [38] ) and especially “ Syrnium huhula ” (= Strix huhula ) or “ Syrnium suinda ” (= Asio flammeus suinda ).
516. Strigidae indet. 2
Quaternary – MG
Syrnium ? sp. e majoribus – Winge 1887: 38.
“duas especies de Syrnium , das quaes uma maior, outra menor” [in part] – Goeldi 1894: 76.
Syrnium sp. (“Mehrere Formen”) [in part] – Lambrecht 1933: 771.
Syrnium sp. [in part] – Mones 1986: 96.
Strigidae indet. 2 – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported a humerus from an unknown origin which he defined as “ Syrnium ? sp.”, larger and slightly different than the record above. He noted it is somewhat larger and considerably different from Asio clamator and slightly larger than in a female “ Syrnium aluco ” (= Strix aluco ).
517. Strigidae indet. 3
Late Holocene – RJ
Strigidae – Carvalho 1984: 117.
Carvalho ( 1984 ) reported a worked strigid femur among the material discovered in 1978 in the Corondó site (RJ-JC-64) in São Pedro da Aldeia, Rio de Janeiro.
518. Strigidae indet. 4
Guérin et al. ( 1996 ) reported material attributable to Asio clamator or Strix huhula from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material. This is one of the taxa the authors highlighted as no longer living in the region today.
Megascops Kaup
519. Megascops choliba (Vieillot)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least two adults and one young individual are present in the material.
Strix Linnaeus
520. Strix virgata (Cassin)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material. This is one of the taxa the authors highlighted as no longer living in the region today.
Glaucidium Boie
521. Glaucidium minutissimum (Wied)
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material. This is one of the taxa the authors highlighted as no longer living in the region today.
522. Glaucidium brasilianum (Gmelin)
Quaternary – MG, PI
Scops brasilianus – Winge 1887: 37.
Scops brasiliensis – Goeldi 1894: 76.
Scops brasilianus – Lambrecht 1933: 771.
Glaucidium brasilianum [in part] – Brodkorb 1971: 225.
Glaucidium brasilianum [ Scops brasilianus ] – Mones 1986: 96.
Glaucidium brasilianum [in part?] – Cuello 1988: 56.
Glaucidium brasilianum [in part] – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, represented by the scapula, coracoid, humerus, ulna, radius, carpometacarpus, femur, tibiotarsus, and tarsometatarsus, apparently with individual variation. Additionally, there is an ulna and a tarsometatarsus from a “saltpeter cave near Escrivânia”, two coracoids and a tarsometatarsus from Lapa da Lagoa do Sumidouro, a humerus and a tibiotarsus from “various caves” [26] , a sternum, a scapula, and a femur of unknown origin, and a scapula, a humerus, and a tarsometatarsus of recent age [28] . He noted that only the distal-most limb bones were determined by comparison with recent bones. The others have the same size and matching characters, which are quite different from Athene cunicularia .
Glaucidium ferox (et aff.) – Winge 1887: 38.
Glaucidium ferox – Goeldi 1894: 76.
Glaucidium ferox (?) – Lambrecht 1933: 771.
Glaucidium brasilianum [in part] – Brodkorb 1971: 225.
Glaucidium ferox – Mones 1986: 96.
Glaucidium brasilianum [in part?] – Cuello 1988: 56.
Glaucidium brasilianum [in part] – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) described remains other than the record above as “ Glaucidium ferox (et aff.)”. They consist of bones of several individuals from Lapa da Escrivânia V, represented by the scapula, coracoid, humerus, ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus. Most bones are slightly smaller than those observed in two recent skeletons. However, two left humeri are considerably smaller and slightly different, and Winge speculated they might belong to Glaucidium pumilum (= minutissimum ), not known from the area. Additionally, there are the two coracoids of a very young individual from Lapa da Escrivânia III.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Athene Boie
523. Athene cunicularia (Molina)
Quaternary – MG, RS
Athene ( Speotyto ) cunicularia – Winge 1887: 38.
Athene cunicularia – Goeldi 1894: 76.
Speotyto cunicularia – Lambrecht 1933: 771.
Speotyto cunicularia – Brodkorb 1971: 226.
Athene ( Speotyto ) cunicularia – Mones 1986: 95.
Athene cunicularia – Cuello 1988: 56.
Athene cunicularia – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) reported two femora, one proximal and one distal end, from Lapa da Escrivânia V, which match this species very well.
Speotyto cunicularia – Rosa 2006b: 232.
Rosa ( 2006b ) reported a bone from level 4 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Speotyto cunicularia – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported this species from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Asio Brisson
524. Asio stygius (Wagler)
Quaternary – MG
Nyctalops stygius – Winge 1887: 37.
Nyctalops stygius – Goeldi 1894: 76.
Nyctalops stygius – Lambrecht 1933: 773.
Asio stygius – Brodkorb 1971: 227.
Asio stygius [ Nyctalops stygius ] – Mones 1986: 95–96.
Asio stygius – Cuello 1988: 56.
Asio stygius – Nascimento and Silveira 2020: 493.
Winge ( 1887 ) associated several bones with at least two individuals from Lapa da Escrivânia V. They are represented by the humerus (proximal and distal ends), carpometacarpus, femur, tibiotarsus (distal end), tarsometatarsus (almost complete), and the first phalanx of the hallux. He noted that only the humeri and femora were determined by comparison, since he had in hands only the skeleton of the torso and the femora and the proximal ends of the humeri of a taxidermied specimen from Lagoa Santa. However, he was almost sure that the other bones belong to the same species and noted that Asio clamator , of which two skeletons were available, is quite distinct.
Trogoniformes AOU
Trogonidae Lesson
Trogon Brisson
525. Trogon sp.
Winge ( 1887 ) reported the distal end of a tibiotarsus from an unknown locality, possibly from Lapa da Escrivânia V. He noted it is much larger than in Trogon surrucura aurantius and somewhat larger than in Trogon viridis , from which it also differs slightly in shape.
526. Trogon surrucura Vieillot
Quaternary – MG
Trogon aff. violaceo – Lund (in Winge 1887 ): 44.
Trogon aurantius – Winge 1887: 44.
[ Trogon ] aurantius – Goeldi 1894: 248.
Trogon aurantius – Lambrecht 1933: 774.
Trogon surrucura – Brodkorb 1971: 247.
Trogon surrucua [ Trogon aurantius ] – Mones 1986: 97.
Trogon surrucua – Cuello 1988: 58.
Trogon surrucura aurantius – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a humerus (“ Trogon aff. violaceo ” in Lund’s catalog) from “a cave near Sumidouro” (other than Lapa da Lagoa do Sumidouro). It corresponds well to that of several specimens of Trogon surrucura aurantius and is considerably smaller than in Trogon viridis .
It was erroneously attributed to Lapa da Lagoa do Sumidouro by Brodkorb ( 1971 ) and Sick ( 1984a , 1993 , 1997 ). This is also present in Lambrecht ( 1933 ) and Cuello ( 1988 ).
Coraciiformes Forbes
Momotidae Gray
Baryphthengus Cabanis & Heine
527. Baryphthengus ruficapillus (Vieillot)
Quaternary – MG
Alcedo aff. amazonæ [in part] – Lund (in Winge 1887 ): 45.
Momotus sp. (næppe M. ruficapillus ) [in part] – Winge 1887: 11.
Momotus ( Baryphthongus ) ruficapillus (et aff.) – Winge 1887: 44–45.
[ Momotus ] ruficapillus – Goeldi 1894: 248.
Momotus ( Baryphthongus ) ruficapillus – Lambrecht 1933: 774.
Baryphthengus ruficapillus – Storer 1960: 79.
Baryphthengus ruficapillus – Fisher 1967: 751.
Baryphthengus ruficapillus – Brodkorb 1971: 254.
Baryphthengus ruficapillus – Mones 1986: 97.
Baryphthengus ruficapillus – Cuello 1988: 59.
Baryphthengus ruficapillus – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported an ulna and the distal end of a tibiotarsus from Lapa da Escrivânia V, a humerus from Lapa da Escrivânia XI, a humerus of an old individual and the coracoid of a young one from Lapa do Marinho II, two ulnae of two individuals from “various caves” [26] , several bones of unknown origin, fragments of recent age [28] , and the proximal end of a humerus from “a cave near Sumidouro” (other than Lapa da Lagoa do Sumidouro). The last one was determined as “ Alcedo aff. amazonæ ” in Lund’s catalog and is very similar to that in Chloroceryle amazona . The total material is represented by the maxilla, coracoid, sternum, humerus, ulna, pelvis, femur, tibiotarsus, and tarsometatarsus. Winge also reported a carpometacarpus from Lapa do Capão Seco, which may belong to another species.
The locality “a cave near Sumidouro” was erroneously presented as Lapa da Lagoa do Sumidouro by Brodkorb ( 1971 ), Cuello ( 1988 ), and Sick ( 1984a , 1993 , 1997 ).
Momotus Brisson
528. Momotus momota (Linnaeus)
Holocene – MT
Momotus momota – Figuti 2005: 156–157, 1.
Figuti ( 2005 ) reported a mummified, incomplete nestling from the uppermost layer of the Santa Elina rock shelter in Jangada, Mato Grosso.
Alcedinidae Rafinesque
Chloroceryle Kaup
529. Chloroceryle amazona (Latham)
Quaternary – MG
Ceryle amazona – Winge 1887: 44.
Ceryle amazona – Goeldi 1894: 248.
Ceryle amazona – Lambrecht 1933: 774.
Chloroceryle amazona – Brodkorb 1971: 253.
Chloroceryle amazona [ Ceryle amazona ] – Mones 1986: 97.
Chloroceryle amazona – Cuello 1988: 59.
Chloroceryle amazona – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a rather incomplete but well-determined humerus from Lapa da Escrivânia V.
Galbuliformes Fürbringer
Bucconidae Horsfield
Malacoptila Gray
530. Malacoptila striata (Spix)
Quaternary – MG
Malacoptila torqvata – Winge 1887: 45.
Monasa torquata – Goeldi 1894: 247.
Malacoptila torquata – Lambrecht 1933: 775.
Malacoptila torquata (= M. striata ) – Storer 1960: 80.
Malacoptila striata – Fisher 1967: 752.
Malacoptila striata – Brodkorb 1971: 257.
Malacoptila striata [ Malacoptila torqvata ] – Mones 1986: 97.
Malacoptila striata – Cuello 1988: 60.
Malacoptila striata – Unwin 1993: 730.
Malacoptila striata – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a humerus from Lapa da Escrivânia V.
Nystalus Cabanis & Heine
531. Nystalus chacuru (Vieillot)
Quaternary – MG
Capito melanotis [in part] – Lund 1841d: 18.
Capito [in part] – Giebel 1847: 20.
Capito [in part] – Pictet 1853: 414.
“un Tamatia ( Capito de Temminck)” [in part] – Liais 1872: 303.
Bucco chacuru – Winge 1887: 45–46.
Bucco chacurú [sic] – Goeldi 1894: 247–248.
Capito sp. (“angeblich aus den Knochenhöhlen Brasiliens”) [in part] – Lambrecht 1933: 769.
Bucco chacuru – Lambrecht 1933: 775.
Bucco (= Nystalus ) chacuru – Storer 1960: 80.
Nystalus chacuru – Fisher 1967: 752.
“ Capito Vieillot 1816 sp., prob. neospecies C. niger (P.L.S. Müller 1776)” – Fisher 1967: 752.
Nystalus chacuru – Brodkorb 1971: 257.
Nystalus chacuru [ Bucco chacuru ] – Mones 1986: 97.
Nystalus chacuru – Cuello 1988: 59.
Nystalus chacuru – Unwin 1993: 730.
Nystalus chacuru – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, represented by the mandible, scapula, coracoid, humerus (in large number), ulna, radius, carpometacarpus, femur, tibiotarsus, and tarsometatarsus. Additionally, there is a mandible from Lapa da Escrivânia III, two humeri (one left and one right) from a “saltpeter cave near Escrivânia”, some bones of unknown origin, and bones of several individuals of recent age [28] , of which Lund determined a humerus. Winge also noted it is easily distinguishable from Malacoptila striata by the humerus and other bones.
Fisher ( 1967 ) attributed a possible record of the capitonid Capito niger from Curvelo or Lagoa Santa based on Capito melanotis of Lund ( 1841d ). However, C. melanotis is synonymous with Nystalus chacuru and, therefore, not a capitonid like C. niger .
?Piciformes Meyer & Wolf
† Gracilitarsidae Mayr
†Eutreptodactylus Baird & Vickers-Rich
532. † Eutreptodactylus itaboraiensis Baird & Vickers-Rich
Early Eocene – RJ
Fig. 31 B
Type locality: São José de Itaboraí, Brazil.
Etymology: Eutreptodactylus , Greek for “easily changeable toe”, and itaboraiensis , referring to the São José de Itaboraí Basin.
“Aves, undetermined (small birds)” [in part?] – Rich 1979: 323, 324.
Eutreptodactylus itaboraiensis sp. nov. [new genus and species] – Baird and Vickers-Rich 1997: 123–127, figs 1–2.
Eutreptodactylus itaboraiensis – Bergqvist et al. 2006: 55.
Baird and Vickers-Rich ( 1997 ) described this new genus and species from material collected by Ney Vidal in 1950 in the São José de Itaboraí Basin. Small avian fossils in this locality had been mentioned by Rich eighteen years before this study (Rich 1979 ).
The material is an incomplete and slightly abraded distal fragment of a right tarsometatarsus (MN 4083-V) with all three trochleae damaged. It was subsequently lost after the study, but photographs, figures, and a cast of the original specimen were housed in the MN collection. The fossil suggests facultative zygodactyly or semizygodactyly (Mayr 2009 ). Mayr et al. ( 2011a ) reported the distal end of a left tibiotarsus (MN 4119-V) from the same locality that may belong to this species (see Neornithes indet. 4).
The taxon was initially attributed to the Cuculidae and then considered its oldest and most primitive representative, possibly belonging to a new subfamily. However, Baird and Vickers-Rich also noted that without the original fossil, more material is necessary before any higher taxon can be confidently erected. Mourer-Chauviré ( 1999 ) found it quite similar to the genus Sylphornis , from the middle Eocene of France, suggesting it should be included in the Sylphornithidae. Mayr ( 2001 ) contested the attribution to the Cuculidae and found similarities between E. itaboraiensis and Gracilitarsus mirabilis (Gracilitarsidae), of the middle Eocene of Messel deposits in Germany, and raised the possibility that they are close relatives. Later, Mayr ( 2005 ) included E. itaboraiensis and Neanis schucherti from the early Eocene of the Green River Formation (USA) in the Gracilitarsidae. He presented a phylogenetic analysis that resulted in a clade composed of Gracilitarsidae, Sylphornithidae, and the Piciformes crown group, but later reiterated (Mayr 2009 , 2016 , 2022 ) that this hypothesis is not yet well established.
Piciformes Meyer & Wolf
Ramphastidae Vigors
533. Ramphastidae indet. 1
Jacobus and Rosa ( 2013 ) reported ramphastid remains from the Deobaldino (RS-S-395) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul. Previously, this record was briefly mentioned as Ramphastos dicolorus by Dias ( 2003 , 2004a ).
534. Ramphastidae indet. 2
Holocene – RS
Ramphastidae – Jacobus 2004: 99.
Jacobus ( 2004 ) reported ramphastid remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
Ramphastos Linnaeus
535. Ramphastos toco Statius Müller
Quaternary – MG
Rhamphastus toco [sic] – Winge 1887: 46.
[ Rhamphastos ] toco [sic] – Goeldi 1894: 247.
Rhamphastus toco [sic] – Lambrecht 1933: 774.
Ramphastos toco – Storer 1960: 81.
“ Ramphastos Linnaeus 1758, 2 spp.” [in part] – Fisher 1967: 752.
Ramphastos toco – Brodkorb 1971: 260.
Ramphastos toco – Sick 1984a: 406.
Ramphastos toco – Mones 1986: 98.
Ramphastos toco – Cuello 1988: 60.
Ramphastos toco – Sick 1993: 372.
[ Ramphastos ] toco – Unwin 1993: 730.
Ramphastos toco – Sick 1997: 492.
Ramphastos toco – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported the proximal part of an ulna and the distal part of a tarsometatarsus from Lapa da Escrivânia V. He noted that only the tarsometatarsus was determined by comparison with recent bones of the same species. The ulna has a similar size.
536. Ramphastos cf. dicolorus Linnaeus
Quaternary – MG
Rhamphastus discolor vel sp. aff. [sic] – Winge 1887: 46.
[ Rhamphastos ] discolorus [sic] – Goeldi 1894: 247.
Rhamphastus discolor [sic] – Lambrecht 1933: 774.
R. discolor [sic] – Storer 1960: 81.
“ Ramphastos Linnaeus 1758, 2 spp.” [in part] – Fisher 1967: 752.
Ramphastos dicolurus [sic] – Brodkorb 1971: 260.
Ramphastos dicolorus [ Ramphastos discolor ] – Mones 1986: 98.
Rhamphastos dicolorus [sic] – Cuello 1988: 60.
Ramphastos dicolurus [sic] – Unwin 1993: 730.
Ramphastos cf. dicolorus – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported an ulna from “various caves” [26] , slightly larger than in Ramphastos dicolorus . There is another ulna from an unknown locality, slightly smaller and insignificantly different than in R. dicolorus and quite different from that of Pteroglossus aracari wiedii and Selenidera maculirostris .
Brodkorb ( 1971 ) erroneously listed it coming from “Lapa da Escrivania?”, which was followed by Cuello ( 1988 ).
Picidae Leach
537. cf. Picidae
Early Holocene – RS
cf. Picidae – Rosa 2009: 151.
Rosa ( 2009 ) reported a possible picid bone from square D6 of the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. The material is deposited in the CEPA collection.
538. Picidae indet. 1
Quaternary – MG
G. sp. indet., forte Leuconerpes candidus – Winge 1887: 46–47.
“talvez coincida com Leuconerpes candidus ” – Goeldi 1894: 247.
cfr. Leuconerpes candidus – Lambrecht 1933: 775.
Leuconerpes candidus – Brodkorb 1971: 265.
cf. Leuconerpes candidus – Mones 1986: 98.
Melanerpes candidus – Cuello 1988: 61.
Picidae indet. 1 – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a humerus from Lapa da Escrivânia V. It is shorter and relatively more robust than in Colaptes campestris , slightly longer and more robust than in Colaptes melanochloros , shorter and more robust than in Celeus flavescens , and much smaller than in Dryocopus lineatus and Campephilus robustus . Winge speculated it might be Melanerpes candidus , although he only had the distal end of a recent humerus available for comparison, which is much more similar to the fossil than the other species in the region. Still, there is a slight difference in shape (perhaps individual variation), and the recent bone is somewhat smaller.
539. Picidae indet. 2
Winge ( 1887 ) reported a coracoid of recent age without attribution of locality. It is of a species smaller than Veniliornis maculifrons[30] , perhaps Veniliornis mixtus cancellatus .
540. Picidae indet. 3
Guérin et al. ( 1996 ) reported material attributable to Dryocopus lineatus or Campephilus melanoleucos from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
541. Picidae indet. 4
Early Holocene – GO
Picidae – Rosa 2004: 256.
Rosa ( 2004 ) reported a bone from the Paranaíba phase cut 2 of the GO-JA-14 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
Picinae Leach
Melanerpes Swainson
542. Melanerpes flavifrons (Vieillot)
Quaternary – MG
Melanerpes flavifrons – Winge 1887: 47.
Melanerpes flavifrons – Goeldi 1894: 247.
Melanerpes flavifrons – Lambrecht 1933: 775.
Melanerpes flavifrons – Brodkorb 1971: 265.
Melanerpes flavifrons – Mones 1986: 98.
Melanerpes flavifrons – Cuello 1988: 60.
Melanerpes flavifrons – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a humerus from Lapa da Escrivânia V, which matches this species well.
Veniliornis Bonaparte
543. Veniliornis maculifrons (Spix)
Quaternary – MG
Picus ( Campias ) maculifrons – Winge 1887: 47.
Campias maculifrons – Goeldi 1894: 247.
Picus ( Campias ) maculifrons – Lambrecht 1933: 775.
Veniliornis maculifrons – Brodkorb 1971: 265.
Veniliornis maculifrons [ Picus ( Campias ) maculifrons ] – Mones 1986: 98.
Veniliornis maculifrons – Cuello 1988: 61.
Veniliornis maculifrons – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported three humeri (one slightly smaller) from Lapa da Escrivânia V, which matches this species well.
Colaptes Vigors
544. Colaptes sp.
Rosa ( 2004 ) reported a bone from the Paranaíba phase square 16H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
545. Colaptes melanochloros (Gmelin)
Quaternary – MG, PI
Chrysoptilus chlorozostus – Winge 1887: 46.
Chrysoptilus chlorozostus – Goeldi 1894: 247.
Chrysoptilus chlorozostus – Lambrecht 1933: 775.
Chrysoptilus melanochloros – Brodkorb 1971: 264.
Chrysoptilus melanochloros [ Chrysoptilus chlorozostus ] – Mones 1986: 98.
Colaptes melanochloros – Cuello 1988: 61.
Colaptes melanochloros – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a femur from Lapa da Escrivânia V. It fits well this species and is quite different in size from the other species available for comparison. The same applies to an incomplete quadrate and the proximal end of a scapula.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
546. Colaptes cf. campestris (Vieillot)
Rosa ( 2001 , updated in 2006b ) reported a bone from level 2 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
547. Colaptes campestris (Vieillot)
Quaternary – MG
Colaptes campester [sic] – Winge 1887: 46.
Colaptes campestris – Goeldi 1894: 247.
Colaptes campester [sic] – Lambrecht 1933: 775.
Colaptes campestris – Brodkorb 1971: 264.
Colaptes campestris – Mones 1986: 98.
Colaptes campestris – Cuello 1988: 61.
Colaptes campestris – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, a humerus from Lapa da Escrivânia XI, and an ulna from “various caves” [26] . The total material is represented by the scapula, coracoid, humerus (in large numbers), ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus. He noted marked differences between these bones and that they are also quite distinct from Campephilus robustus , Dryocopus lineatus , Colaptes melanochloros , and Celeus flavescens . However, some slightly smaller tarsometatarsi may belong to Melanerpes candidus .
Cariamiformes Fürbringer
†Paleopsilopterus Alvarenga
548. † Paleopsilopterus itaboraiensis Alvarenga
Early Eocene – RJ
Fig. 31 D
Type locality: Holotype collected in the limestone basin of the São José village, district of Cabuçu, municipality of Itaboraí, state of Rio de Janeiro, Brazil, among the marl that fills the limestone galleries.
Etymology: Paleopsilopterus , with the Greek “ palaeos ” (ancient), based on the hypothesis that it was an older and perhaps ancestral form of Psilopterus , and itaboraiensis, referring to the São José de Itaboraí Basin.
Aves [“Ainda por determinar”; in part?] – Paula Couto 1958a: 11.
Aves [“Diversas, não identificadas”; in part?] – Paula Couto 1970: 909.
Aves [“fragmentos de ossos longos”; in part?] – Palma 1973: 40.
Aves [“fragmentos de ossos longos”; in part?] – Palma and Brito 1974: 400.
“restos de diversas aves, metatarsais, vértebras, pré-maxilas, etc.” [in part?] – Price (in lit. 1977 in Sick 1984a ): 65.
Aves [“representadas por ossos longos”; in part?] – Francisco and Souza Cunha 1978: 390, 402, 408.
Psilopteridae – Alvarenga 1983b: 6.
Paleopsilopterus itaboraiensis sp. n. [new genus and species] – Alvarenga 1985a: 17–20, figs 1–4.
Palaeopsilopterus itaboraiensis [sic] – Mones 1986: 89.
Paleopsilopterus itaboraiensis – Cuello 1988: 18.
“fragmentos de ossos longos de aves” [in part?] – Brito 1989: 60.
Paleopsilopterus itaboraiensis – Alvarenga 1993a: 62.
Paleopsilopterus itaboraiensis – Unwin 1993: 725.
Paleopsilopterus itaboraiensis – Alvarenga 1997: 123.
Paleopsilopterus itaboraiensis – Kellner 1998: 654, 659.
Paleopsilopterus itaboraiensis – Alvarenga 1999b: 58–59, 39C.
Paleopsilopterus – Alvarenga and Höfling 2000: 590.
Paleopsilopterus itaboraiensis – Alvarenga and Höfling 2003: 84–85, figs 29, 34.
Paleopsilopterus – Alvarenga and Höfling 2004: 828.
Paleopsilopterus itaboraiensis – Bergqvist et al. 2006: 55, 61.
Paleopsilopterus itaboraiensis – Agnolín 2009b: 21, 18.
Paleopsilopterus itaboraiensis [in part] – Alvarenga et al. 2011: 191–192, 7.2.
Paleopsilopterus – Alvarenga and Höfling 2011: 128.
Alvarenga ( 1985a ) described this new genus and species from remains collected in the late 1940s in the São José de Itaboraí Basin. They were briefly alluded to by him previously (Alvarenga, 1983b ).
The holotype is the proximal end of a right tarsometatarsus with the upper extremity of the hypotarsus partially damaged (MN 4040-V), collected by Ney Vidal in 1948. The paratypes are the two tibiotarsi of a single individual (MCT.R.1431), incomplete at the proximal end, very deformed and encrusted with limestone, collected by Júlio da Silva Carvalho in 1949. Both were restored. Although the condyles of the right tibiotarsus articulate rightly with the holotype tarsometatarsus, they likely belong to different individuals, due to the different collection dates and state of conservation, besides the lack of more detailed information on the collection site. Its body mass was estimated at 4.198 kg (Taranto and Bergqvist 2010 ).
Its systematic placement was widely debated. Alvarenga (1983b, 1985a ) initially included it in the Psilopteridae, representing then the first record of this family outside Argentina. In favor of a position close to the Phorusrhacidae (possibly even much closer to a common ancestor of the phorusrhacoids), he highlighted the greater size and robustness of the bones when compared with Psilopterus (middle Oligocene to the late Miocene of Argentina), and the close similarity to the tarsometatarsus of Paraphysornis brasiliensis , resembling a miniaturized version of it. Later, Alvarenga and Höfling ( 2003 ) included it in the Psilopterinae subfamily within Phorusrhacidae and highlighted the similarity with Procariama simplex (late Miocene and early Pliocene of Argentina). Mayr ( 2009 , 2022 ) commented that the assignment to that subfamily was based on general similarities but should be verified with additional specimens and supported by derived characteristics shared between the involved taxa. Agnolín ( 2009b ) excluded the taxon from the Phorusrhacoidea in favor of a placement with the European Cariamoidea Idiornithidae. Tambussi and Degrange ( 2013 ) also disagreed with the placement within the Phorusrhacidae Psilopterinae. However, not convinced of the arguments used by Agnolín ( 2009b ) for its inclusion in the Idiornithidae, they considered it as Cariamiformes incerti familiae due to its fragmentary nature and a set of characters it shares with other Cariamiformes. Alvarenga et al. ( 2011 ) kept the tentative classification within the Phorusrhacidae Psilopterinae following the characteristics indicated in Alvarenga and Höfling ( 2003 ). Agnolín ( 2013 ), rebutting Alvarenga et al., kept his arguments previously presented (Agnolín 2009b ). Mayr ( 2016 ) pointed out that additional specimens are needed to clarify its classification.
Paleopsilopterus itaboraiensis was considered the oldest known member of the Phorusrhacidae (Alvarenga 1985a , Alvarenga and Höfling 2003 , Mayr 2009 , Alvarenga et al. 2011 , Mayr 2016 ). If excluded from Phorusrhacoidea, the best-documented Phorusrhacidae record in South America would come from the late Eocene of Chubut, Argentina (Tonni and Tambussi 1986 apud Agnolín 2009b ). Additionally, material from a slightly older age (middle Eocene) from the Guabirotuba Formation in Paraná is awaiting a more detailed description (see Phorusrhacidae indet. 1; Angst and Buffetaut 2017 ).
The proximal end of a tarsometatarsus from the late Eocene of the Sarmiento Formation in Gran Hondonada (province of Chubut, Argentina) was assigned to the genus Paleopsilopterus by Acosta Hospitaleche and Tambussi ( 2005 ). Agnolín ( 2009b ) considered it a member of the Idiornithidae, representing a new genus and species. Tambussi and Degrange ( 2013 ), mentioning Degrange (2012 apud Tambussi and Degrange 2013 ), who indicated the material is not a Phorusrhacidae, noted that it is not possible to verify its systematic position without any additional material.
Phorusrhacidae – Taranto and Bergqvist 2009: 51R.
Phorusrhacidae – Taranto et al. 2009a: 287.
Paleopsilopterus itaboraiensis – Taranto and Bergqvist 2010: 118.
Paleopsilopterus itaboraiensis [in part?] – Metello et al. 2014: 84.
Paleopsilopterus itaboraiensis [in part?] – Metello and Bergqvist 2014: 151.
Taranto and Bergqvist ( 2009 ) and Taranto et al. ( 2009a ) associated with the Phorusrhacidae an ungual phalanx (UFRJ 03-AV) from the São José de Itaboraí Basin. It was later associated with Paleopsilopterus itaboraiensis by Taranto and Bergqvist ( 2010 ). Metello and Bergqvist ( 2014 ) mentioned two ungual phalanges associated with this species, possibly listing this material as MCT 1837-R. It figures among material long stored in the Departamento de Geologia of UFRJ and has approximately half the length of that of Phorusrhacos longissimus .
Paleopsilopterus itaboraiensis [in part] – Alvarenga et al. 2011: 191–192, 7.2.
Alvarenga et al. ( 2011 ) tentatively attributed to this species five ungual phalanges collected by Rubens da Silva Santos during the 1970s in the same locality as the type material. They are assigned to digits I (MHNT-VT-5320), II (MHNT-VT-5316), III (MHNT-VT-5317 and MHNT-VT-5319), and IV (MHNT-VT-5318).
Metello et al. ( 2012a ) and Metello and Bergqvist ( 2014 ) associated with this species another ungual phalanx (MCT 1836-R), attributed to digit IV. Metello et al. ( 2012a ) strangely labeled it as the first avian ungual phalanx described for the Paleocene (=early Eocene following Gelfo et al. 2009 ) of South America. Assessing the curvature of the phalanges, Metello and Bergqvist ( 2014 ) found support for the assumption of terrestrial habits, reinforcing the theory that the Phorusrhacidae attacked their prey with their massive skull, but not excluding the possibility of use in their raptorial habits.
†Itaboravis Mayr, Alvarenga & Clarke
549. † Itaboravis elaphrocnemoides Mayr, Alvarenga & Clarke
Early Eocene – RJ
Fig. 31 C
Type locality: São José de Itaboraí, Rio de Janeiro, southeastern Brazil
Etymology: Itaboravis , with the Latin avis , “bird of Itaboraí”, and elaphrocnemoides , with the Latinized suffix oideus , referring to the similarities with the genus Elaphrocnemus .
“Une forme très proche du genre Elaphrocnemus ” [in part?] – Alvarenga (in Mourer-Chauviré 1999 ): 87.
“idiornithid-like birds” [in part?] – Mayr 2009: 142.
“Idiornithidae indeterminado” [in part?] – Agnolín 2009b: 20.
“unpublished Cariamae, closely related to the European Idiornithidae” [in part?] – Alvarenga et al. 2011: 201.
Itaboravis elaphrocnemoides , gen. et sp. nov. [new genus and species]- Mayr et al. 2011a: 680, 1A, G–H.
Itaboraves elaphrocnemoides [lapsus] – Taranto 2012: 11, 51.
Mayr et al. ( 2011a ) described this new genus and species from remains found in the São José de Itaboraí Basin. Mourer-Chauviré ( 1999 ), through information provided by Herculano Alvarenga, already pointed out the presence of this taxon with close similarity to Elaphrocnemus , a European genus of the late Eocene and Oligocene with three known species (Mayr 2016 ).
The remains consist of a left coracoid (MN 4114-V, holotype), a right humerus (MN 4113-V), and the distal end of a left humerus (MN 4121-V). Alvarenga, one of the authors of the study, was informed by Fausto Luiz de Souza Cunha in 1985 that specimens MN 4113-V and MN 4114-V stem from the same calcareous matrix block of about 1.4 kg, which also yielded many small, non-avian bones. Furthermore, both bones share the same coloration, match in size, and are similar to those of Elaphrocnemus , which led the authors to believe they belonged to the same individual. The coracoid is from a bird the size of Crypturellus tataupa and most closely resembles that of Elaphrocnemus . The humerus is similar to that of Elaphrocnemus but also presents certain affinities to the Tinamidae, and indicates rather weak flight capacities, perhaps comparable to those of living tinamids.
Mayr et al. considered the affinities with Elaphrocnemus best supported by current evidence and tentatively attributed the Itaboraí material to the Cariamae (sensu Mayr 2009 ). They noted, however, that this classification is mainly based on overall similarity, and, if the indeterminate carpometacarpus MN 4115-V (see Neornithes indet. 3) indeed belongs to this taxon, Itaboravis may be a stem group representative of the Tinamidae with a very different coracoid morphology.
Elaphrocnemus was for a long time (including the time of Itaboravis description) classified within the Idiornithidae, a stem group of European Cariamiformes. Its phylogenetic affinities are not convincingly resolved, presenting similarities with both Cariamiformes and Opisthocomiformes, and, in any case, it likely does not belong to a clade comprising the Idiornithidae, Phorusrhacidae, and Cariamidae (Mayr 2016 ). Accordingly, Mayr ( 2016 ) also noted that the placement of Itaboravis is equally uncertain. Regardless of its classification, I. elaphrocnemoides supports the evidence that the earliest Cenozoic avifaunas were already diverse.
Cariamidae Bonaparte
Cariama Brisson
550. cf. Cariama cristata (Linnaeus)
Pacheco et al. ( 2007 ) reported avian remains including a specimen that possibly belonged to this species from the Maracaju 1 site in Maracaju, Mato Grosso do Sul. Although Pacheco et al. mentioned a single bone from the site’s layer IV, Pacheco and Martins ( 2009b ) mentioned both a “cariamid” bone from layer I and a “seriema” bone from layer IV.
551. Cariama cristata (Linnaeus)
Quaternary – GO, MT, MG, PI, SP
Dicholophus cristatus – Winge 1887: 31.
Dicholophus cristatus – Goeldi 1894: 559.
Cariama cristata – Lambrecht 1933: 761.
Cariama cristata – Brodkorb 1967: 173.
Cariama cristata [ Dicholophus cristatus ] – Mones 1986: 89.
Cariama cristata – Cuello 1988: 44.
Cariama cristata – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported part of the proximal end of a slightly rolled tarsometatarsus from Lapa da Lagoa do Sumidouro.
“seriemas” – Schorr 1976: 100.
Schorr ( 1976 ) reported this species among the remains from the GO-JA-01, GO-JA-14, and GO-JA-20 rock shelters in Serranópolis, Goiás.
“siriema” – Alves 2008: 170.
Alves ( 2008 ) reported “seriema” remains from area III (at least one individual) of the Capelinha I site in Cajati, São Paulo, aged 9,250±50 years BP.
Cariamidae – Pacheco 2008: 229.
Pacheco ( 2008 ) reported a fragment of a cariamid proximal tibiotarsus from the Santa Elina rock shelter in Jangada, Mato Grosso.
Cariama cristata – Barbosa 2017: 126.
Barbosa ( 2017 ) reported this species from Toca do Alto da Serra do Capim in Guaribas and Toca dos Coqueiros in Coronel José Dias, Piauí. These holocenic remains were associated with combustion structures (three bones at Toca dos Coqueiros).
† Phorusrhacidae Ameghino
552. † Phorusrhacidae indet. 1
Sedor et al. ( 2014b ) reported phorusrhacid remains from the Guabirotuba Formation ( Fig. 1 .5) in the Curitiba Basin, Paraná, with an estimated age at the end of the middle Eocene (Barrancan SALMA; Sedor et al. 2016 ). The fossils were found on an outcrop at the border of the municipalities of Curitiba and Araucária (Sedor et al. 2014a ). They are slightly older than material from the late Eocene of Chubut, Argentina, reported to be the oldest best-documented Phorusrhacidae record in South America (Angst and Buffetaut 2017 ).
The material consists of a large isolated cervical vertebra, corresponding to the caudalmost part of the neck (C7 to C10), and the distal end of a tarsometatarsus, deposited in the paleontological collection of the Museu de Ciências Naturais da Universidade Federal do Paraná.
The bird lived in floodplains, in predominantly wet climatic conditions alternating with drier periods. The associated paleofauna included gastropods and other invertebrates, bony fishes, anurans, testudines, sebecosuchian crocodilians, cingulates, notoungulates, astrapotheres, and metatherians (Sedor et al. 2014a , 2016 ).
553. † Phorusrhacidae indet. 2
Late Miocene – AC
“grandes aves” – Bocquentin and Souza Filho 1990: 233.
Phorusrhacidae – Alvarenga 1992: 254.
[might be referred to the] Phorusrhacinae – Tambussi and Noriega 1996: 252.
Phorusrhacinae indet. – Latrubesse et al. 1997: 112.
Phorusrhacidae indet – Bocquentin and Silva 1998: 154.
Phorusrhacinae gen. et sp. indet. – Negri and Ferigolo 1999: 18.
“Género y especie indeterminados” – Agnolín 2009b: 56.
Bocquentin and Souza Filho ( 1990 ) first reported the occurrence of large birds from the Solimões Formation of the Niterói site based on a phalanx. Later, Alvarenga ( 1992 ; through personal communication with Kenneth E. Campbell) associated this record with the Phorusrhacidae and mentioned several unstudied bones of a gigantic bird collected by researchers of the LACM. They were later listed by Latrubesse et al. ( 1997 ), Bocquentin and Silva ( 1998 ), Negri and Ferigolo ( 1999 ), and Agnolín ( 2009b ).
The material is composed of pedal phalanges, deposited without number in the LPP collection of UFAC (Bocquentin and Silva 1998 , Agnolín 2009b ). Tambussi and Noriega ( 1996 ) noted that these fragmentary remains might be referred to Phorusrhacinae, which, if confirmed with additional material, would represent the subfamily’s first member from Brazil. Agnolín ( 2009b ) commented that, despite being incomplete, they could be attributed to the subfamily Phorusrhacinae (sensu Agnolín 2009b ) for their large size and for presenting a higher than wide proximal articular face, a character shared with the genera Devincenzia , Paraphysornis , and Physornis .
† Physornithinae Agnolín
†Paraphysornis Alvarenga
554. † Paraphysornis brasiliensis (Alvarenga)
Late Oligocene/Early Miocene – SP
Figs 4A, 29, 32B
Type locality: Municipality of Tremembé, district of Padre Eterno, near the right margin of the Paraíba do Sul river, state of São Paulo.
Etymology: Paraphysornis , with the Greek “ para” (close to), alluding to the related genus Physornis , and brasiliensis , referring to Brazil.
Physornis brasiliensis sp. n. [new species] – Alvarenga 1982: 697–712, figs 1–16.
“ave gigante” – Sick 1984a: 65.
Physornis brasiliensis – Mones 1986: 88.
Physornis brasiliensis – Cuello 1988: 16.
Physornis brasiliensis – Alvarenga 1993a: 62, 34.
Physornis brasiliensis – Alvarenga 1993b: 22, figs 15–16.
Paraphysornis brasiliensis [new genus] – Alvarenga 1993c: 403–406, figs 1–2.
Paraphysornis brasiliensis – Alvarenga 1997: 123, 34.
Paraphysornis brasiliensis – Alvarenga 1999b: 33–34, figs 2, 8B, 10A–B, 12B, 13E, 23B, 24B, 25.
Paraphysornis brasiliensis – Kellner 1998: 654, 659.
Paraphysornis brasiliensis – Alvarenga and Höfling 2000: 590, 31.9.
Paraphysornis brasiliensis – Alvarenga and Höfling 2003: 70, figs 1–4, 6, 15–17, 34.
Paraphysornis brasiliensis – Alvarenga and Höfling 2004: 828, 44.9.
Paraphysornis brasiliensis – Agnolín 2009b: 53–54, figs 1–2, 23, 29, 32, 42, 52.
Paraphysornis brasiliensis – Jones 2010: 10, figs 2.4–2.5, 2.8, 4.4–4.9, E.6–E.7.
Paraphysornis brasiliensis – Alvarenga et al. 2011: 201, figs 7.1, 7.3–7.6, 7.8–7.10.
Paraphysornis brasiliensis – Alvarenga and Höfling 2011: 128, 7.11.
Paraphysornis brasiliensis – Angst and Buffetaut 2017: 137, 5.9.F.
Paraphysornis brasiliensis – Angst and Chinsamy 2017: 17–37, 1B.
Paraphysornis brasiliensis – Carmo et al. 2024: 5, 3A–B.
Paraphysornis brasiliensis (MHNT-VT-5000, restored holotype as displayed in MHNT), highlighted elements: femora, right (A) and left (B); tibiotarsi, right (C) and left (D); fibulae, right (E) and left (F); tarsometatarsi, right (G) and left (H); cervical vertebrae (I); thoracic vertebrae (J); dorsal ribs (K, L, M); sternal ribs (N, O); skull cap fragment (P); left supraorbitary (Q); fragment of the tip of the premaxilla (R); mandible (S); left quadrate (T); left pterygoid (U); left coracoid (V); possible left clavicle (W); humeri, right (X) and left (Y); left ulna (Z); left radius (Aa); carpometacarpi, left (Ab) and right (Ac); proximal phalanx of the left second digit (Ad); proximal phalanx of the left second digit (Ae); caudal segments of the pubes (Af, Ag); cranial fragments of the ischium, left (Ah) and right (Ai); antero-superior part of the pelvis (Aj); caudal vertebrae (Ak); hallux metatarsal (Al); left pedal phalanges (Am) (G. Couto Ribeiro). Scale bar: 100 mm.
Alvarenga ( 1982 ) described this new species from well-preserved and sparse fragments found in the montmorillonite clays of the Tremembé Formation, 2 or 3 m below the pyrobituminous shales. They were collected during several months between 1977 and 1978.
The holotype is an incomplete skeleton (MHNT-VT-5000, originally DGM 1418-R), which lacks most of the skull and premaxilla, pelvis, and sternum, with the other bones well represented and some even complete—about 70% of the skeleton is available (Alvarenga et al. 2011 ). Alvarenga also mentioned many bone fragments accompanying the specimen, which may provide further anatomical data. In the original description, the depicted elements were the mandible lacking the right branch, left quadrate, left pterygoid, left supraorbital, skullcap fragment, premaxilla extremity fragment, cervical vertebrae (C-1 [atlas], C-2 [axis]), C-3, C-10, and C-11, out of a total of thirteen), thoracic vertebrae (T-1, T-2, T-3, T-4, and T-5; a sixth appears to be attached to the pelvis), caudal vertebrae (one more anterior and one more posterior), the anterosuperior portion of the pelvis, fragments of dorsal ribs (seven) and sternal ribs (two fragmented and one complete), left coracoid (which indicates, along with the sternal ribs, the presence of a well-developed sternum), left and right humeri, left ulna, proximal half of the left radius, left and right carpometacarpi, proximal and distal phalanges of the major digit of the left wing, left and right femora (both lacking a segment of the diaphysis), left and right tibiotarsi, left and right fibulae, left and right tarsometatarsi, metatarsal bone of the left hallux, nine phalanges of the left foot (second and ungual of digit II, third and ungual of digit III and first, second, third, fourth, and ungual of digit IV), and the fourth phalanx of digit IV of the right foot. A partial reconstruction of the skeleton with the posture in life and a life reconstruction of the bird were presented. Alvarenga et al. ( 2011 ) depicted a fragment that appears to belong to the cranial end of the left clavicle and fragments of the pelvis [39] that preserved the cranial parts of the ischium and the caudal projections of the two pubes, all not originally described by Alvarenga ( 1982 ). Also depicted are three cervical vertebrae (third and possibly the tenth and eleventh), one thoracic vertebra, the reconstructed first pre-synsacral thoracic vertebrae, and remains of the cranial end of the iliac dorsal crest.
The bird had a large head, with a strong jaw approximately half a meter long. After restoring the skeleton, Alvarenga estimated it reached around 2 m in height and up to 3 m at the tip of the beak when keeping the legs and spine stretched. Later, Alvarenga and Höfling ( 2003 ) estimated 140 cm of height on the back, reaching 240 cm with the head stretched, and a weight of approximately 180 kg, being perhaps the smallest of the Brontornithinae (sensu Alvarenga and Höfling 2003 ). The wings were quite small, and the ulna had no marks of remige insertions. The second toe has a very wide, curved, and strong claw, probably important for holding prey.
Alvarenga ( 1982 ) proposed that, due to the relatively short and thick tarsometatarsus, the bird was apparently slower and unfit to run when compared with other phorusrhacids, perhaps having been a scavenger. Alvarenga and Höfling ( 2003 ) supported this theory of graviportality (and applied it to the other Brontornithinae) through the fact that the lacustrine Tremembé Formation presented periods of drought with a high fish mortality rate, and the only known skeleton of the species may have come from an individual that succumbed into this swampy land while looking for fish and other dead animals. The attribution of necrophagy as the main dietary source of the larger species, while the smaller ones would have been active predators, was also addressed by other authors (Tonni 1977 and Tambussi and Hoffmann 1998 apud Agnolín 2009b ). Jones (2005 apud Jones 2010 ) reported that the tibiotarsus conformation of Paraphysornis (as well as that of Brontornis ) could be associated with the ability to break bones, which was proposed to other species by Blanco and Jones ( 2005 ) and related with inferences about possible scavenging habits.
Jones ( 2010 ) proposed that Paraphysornis and Brontornis could have had horny sheats in their curveless ungual phalanges, especially in digit III, as in large extant cursorial birds (e.g., struthionids, rheids, otidids). However, he also noted (Jones 2005 apud Jones 2010 , Jones 2010 ) that, from the ungual phalanges of the Brontornithinae (sensu Alvarenga and Höfling 2003 ), the graviportal condition can only be expressed due to its robust character and lateral expansion of the pedal phalanges and, added to the study of the proportions of the bones of the legs, they would not have had prominent cursorial habits.
Tambussi and Degrange ( 2013 ) noted that the Paraphysornis tarsometatarsus is overall similar to that of Dromornis stirtoni (late Miocene of Australia) and pointed out the contrast of Alvarenga’s ( 1982 ) theory with that of Murray and Vickers-Rich ( 2004 ), who argued that high masses would not limit the cursorial skills of the dromornithids and even those estimated at 500 kg were able to run. However, the pelvis, an important element for establishing locomotor and postural habits (Degrange 2012 apud Tambussi and Degrange 2013 ), was not found complete. Tambussi and Degrange further noted that the cranial remains of Paraphysorni s include the mandible and a quadrate bone whose morphology on its own is not indicative of feeding habits, and assignment with any should be taken with caution.
Angst et al. ( 2016 ), Angst and Chinsamy-Turan ( 2016 ), and Angst and Chinsamy ( 2017 ) also proposed that Paraphysornis had a relatively slow graviportal locomotion due to the biomechanical limitations of its size and body mass. However, they noted that, as its skull clearly demonstrates adaptation to a carnivorous diet, it may have relied on a specific hunting method, having been an ambush predator or, as already mentioned in its original description, a scavenger. Angst and Buffetaut ( 2017 ) suggested that the more robust phorusrhacids might have attacked large and slow prey, such as astrapotheres and pyrotheres. LaBarge et al. ( 2024 ) analysis also recovered it as a graviportal, likely ambush predator.
In its original description, the taxon was provisionally attributed to the genus Physornis (originally from the Oligocene of the province of Santa Cruz, Argentina), in the subfamily Brontornithinae. Subsequently, Alvarenga ( 1993c ) examined phorusrhacid material in museums in Argentina, England, France, and the USA, and compared them with the fossils from the Tremembé Formation. He concluded that the Brazilian species differs significantly from Brontornis burmeisteri and Physornis fortis , although it is closer to the latter, and erected the new genus Paraphysornis .
Alvarenga and Höfling ( 2003 ) kept the taxon within the Brontornithinae, as perhaps its smallest representative. Agnolín ( 2007 ) excluded Brontornis from the Phorusrhacidae in favor of a basal position in the Anseriformes and coined the subfamily Physornithinae for Physornis and Paraphysornis . Later, Agnolín ( 2009b ) included Paraphysornis in the Phorusrhacinae Physornitini, along with Physornis and Devincenzia . Alvarenga et al. ( 2011 ), not supporting Agnolín’s hypothesis, kept Paraphysornis along with Brontornis and Physornis in the Phorusrhacidae Brontornithinae, with Devincenzia in the Phorusrhacinae. Angst and Buffetaut ( 2017 ) followed Agnolín for Brontornis and kept Paraphysornis in the Phorusrhacinae. LaBarge et al. ( 2024 ) included it, along with Physornis , in the Phorusrhacidae Physornithinae, and noted that a definitive systematic assessment of Brontornis will require a phylogenetic reappraisal including many more modern and fossil neognaths than their study focused on, in particular Gastornithidae and Dromornithidae.
Falconiformes Bonaparte
555. cf. Falconiformes (sensu lato?) 1
Early Holocene – GO
cf. Falconiformes – Rosa 2004: 237.
Rosa ( 2004 ) reported three bones of at least one individual from the Paranaíba phase square 16H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
556. cf. Falconiformes (sensu lato?) 2
Late Holocene – RS
cf. Falconiformes – Rosa 2006c: 250.
Rosa ( 2006c ) reported two bones from levels 4 and 6 of square 1 of the RS-LC-80 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
557. Falconiformes (sensu lato?) indet. 1 (spp.?)
Rosa ( 2006b ) reported four bones (one from level 1, one from level 2, and two from level 4) from the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. This latter report differs slightly from Rosa ( 2001 ), who mentions two falconiform bones, one from level 2 and one from level 5. The material is deposited in the IAP/Unisinos collection.
558. Falconiformes (sensu lato?) indet. 2 (spp.?)
Rosa ( 2004 ) reported Falconiformes (sensu lato?) remains from the Paranaíba phase of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material consists of five bones of at least one individual from square 12H, two bones of at least one individual from 14H, a bone from 18H, and a bone from 18I, and is deposited in the IAP/Unisinos collection.
559. Falconiformes (sensu lato?) indet. 3
Early Holocene – SP
“Falconiforme” – Alves 2008: 100.
Alves ( 2008 ) reported Falconiformes (sensu lato? As it is also referred to as a “hawk” in the text) remains from area III (at least one individual) of the Capelinha I site in Cajati, São Paulo, aged 9,250±50 years BP.
560. Falconiformes (sensu lato?) indet. 4
Schmitz and Ferrasso ( 2011 ) reported a falconiform humerus from the Guarani site Itapiranga I (SC-U-1) in Itapiranga, Santa Catarina. The material was discovered in January 1957 and is deposited in the IAP/Unisinos collection.
561. Falconiformes (sensu lato?) indet. 5
Silva ( 2013 , 2014 ) reported falconiform material among remains collected by the staff of NEA-UFPE, starting in the 1990s, in rock shelters of the Pedra do Alexandre site in Carnaúba dos Dantas, Rio Grande do Norte.
Falconidae Leach
562. cf. Falconidae
Rosa ( 2009 ) reported eight possible falconid bones (one from square A6 of the site’s third occupation, one from C7, one from D5, four from D6, and one from D7) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. The material is deposited in the CEPA collection.
563. Falconidae indet.
Late Holocene – SC
“Falconídeos” – Schmitz et al. 1992: 110.
Schmitz et al. ( 1992 ) reported falconid remains from the Armação do Sul site in Ilha de Santa Catarina, Florianópolis, Santa Catarina.
Herpetotherinae Lesson
Micrastur Gray
564. Micrastur sp.
Quaternary – MG
Micrastur sp. e majoribus ( M. melanoleucus ?) – Winge 1887: 34.
Falconines [?] [in part] – Goeldi 1894: 76.
M. cfr. melanoleucus (?) – Lambrecht 1933: 751.
Micrastur semitorquatus – Brodkorb 1964: 296.
Micrastur semitorquatus – Mones 1986: 85.
Micrastur semitorquatus – Cuello 1988: 39.
Micrastur sp. – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported the distal end of a humerus and the distal end of a tibiotarsus of unknown origin. They are larger than in Micrastur ruficollis , and the shape matches that of Micrastur semitorquatus . However, they are somewhat smaller than the specimen used for comparison, which is large and probably a female. The fossil material possibly belonged to a small male of that species. Winge also noted they differ considerably from Milvago and Caracara , and that no equivalent material from Herpetotheres was available for comparison.
565. Micrastur ruficollis (Vieillot)
Quaternary – MG
Micrastur ruficollis – Winge 1887: 34.
Falconines [?] [in part] – Goeldi 1894: 76.
Micrastur ruficollis – Lambrecht 1933: 751.
Micrastur ruficollis – Brodkorb 1964: 296.
Micrastur ruficollis – Mones 1986: 85.
Micrastur ruficollis – Cuello 1988: 39.
Micrastur ruficollis – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported two left humeri from Lapa da Escrivânia V.
Caracarinae d’Orbigny
Caracara Merrem
566. Caracara plancus (Miller)
Quaternary – MG, PI, RS
Polyborus tharus – Winge 1887: 34.
Falconines [?] [in part] – Goeldi 1894: 76.
Polyborus tharus – Lambrecht 1933: 750.
Caracara plancus – Brodkorb 1964: 296.
Caracara plancus [ Polyborus tharus ] – Mones 1986: 85.
Polyborus plancus – Cuello 1988: 39.
Caracara plancus – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, most of young or very young birds, represented by the coracoid, humerus, ulna, femur, tibiotarsus, and tarsometatarsus. All are slightly smaller than that observed in two recent skeletons. There is also a tarsometatarsus of a very young individual from Lapa da Escrivânia XI.
Accipitridae [?] [in part] – Guérin et al. 1993a: 198.
Falconidae [?] [in part?] – Guérin et al. 1993a: 198.
Accipitridae [?] [in part] – Guérin et al. 1993b: 328.
Falconidae [?] [in part?] – Guérin et al. 1993b: 328.
Polyborus plancus – Guérin et al. 1996: 84.
Polyborus plancus – Guérin et al. 2002: 136.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
Rosa ( 2006b ) reported three bones (two from level 2 and one from level 4) from the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Polyborus plancus – Silva et al. 2006: 338.
Silva et al. ( 2006 ) reported this species from the RS-RG-49 site in Rio Grande, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Milvago Spix
567. Milvago chimachima (Vieillot)
Quaternary – MG
Milvago chimachima – Winge 1887: 34.
Falconines [?] [in part] – Goeldi 1894: 76.
Milvago chima chima [sic] – Lambrecht 1933: 749.
Milvago chimachima – Brodkorb 1964: 296.
Milvago chimachima – Mones 1986: 85.
Polyborus chimachima – Cuello 1988: 39.
Milvago chimachima – Nascimento and Silveira 2020: 494.
Winge ( 1887 ) reported a tarsometatarsus from Lapa da Escrivânia V. He noted that numerous Milvago bones were found in that cave, including very young specimens, but most could not be determined at the species level.
568. Milvago chimango (Vieillot)
Winge ( 1887 ) reported a tarsometatarsus (well distinguished from Milvago chimachima ) and two tibiotarsi from Lapa da Escrivânia V. Many bones he determined as Milvago might belong to this taxon rather than to M. chimachima , despite M. chimango not being known to live in the Lagoa Santa region.
Falconinae Leach
Falco Linnaeus
569. Falco sp. 1
Chahud ( 2021 ) reported the distal part of a right tarsometatarsus (ST-2054) of an adult individual from Lapa do Santo in Matozinhos, Minas Gerais. The material is deposited in the LEEH-IB-USP collection. Chahud noted that its proportions agree with that of Falco peregrinus , a species with various extant records in the region. It was found at a different point in the site and thus might be of a different age than the tinamid also described by Chahud.
570. Falco sp. 2
Quaternary – MG
Falco sp. – Seersholm et al. 2021: 2067.
Seersholm et al. ( 2021 ) reported this genus among the bones collected by Lund in Lapa da Escrivânia V that were analyzed through bulk bone metabarcoding. It was also detected morphologically in one of the studied samples.
571. Falco sparverius Linnaeus
Quaternary – MG, PI
Falco ( Tinnunculus ) sparverius – Winge 1887: 34–35.
Falconines [?] [in part] – Goeldi 1894: 76.
Falco sparverius – Lambrecht 1933: 750.
Falco sparverius – Brodkorb 1964: 296.
Falco sparverius – Mones 1986: 85.
Falco sparverius – Cuello 1988: 40.
Falco sparverius – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported remains of several individuals (both adult and young) from Lapa da Escrivânia V, a coracoid from Lapa da Escrivânia XI, and several bones of recent age [28] , two humeri of which were determined by Lund. The total material is represented by the thoracic vertebra, scapula, coracoid, sternum, humerus (in large numbers), ulna, carpometacarpus, pelvis, femur, tibiotarsus, and tarsometatarsus. Markedly individual variation exists, especially in the material from Lapa da Escrivânia V, with individuals larger than a female Falco sparverius , but it was not possible to establish a limit between smaller and larger sets of bones.
Accipitridae [?] [in part] – Guérin et al. 1993a: 198.
Falconidae [?] [in part?] – Guérin et al. 1993a: 198.
Accipitridae [?] [in part] – Guérin et al. 1993b: 328.
Falconidae [?] [in part?] – Guérin et al. 1993b: 328.
Falco sparverius – Guérin et al. 1996: 84.
F. sparverius – Guérin et al. 2002: 136.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material.
572. Falco rufigularis Daudin
Quaternary – PI
Accipitridae [?] [in part] – Guérin et al. 1993a: 198.
Falconidae [?] [in part?] – Guérin et al. 1993a: 198.
Accipitridae [?] [in part] – Guérin et al. 1993b: 328.
Falconidae [?] [in part?] – Guérin et al. 1993b: 328.
Falco rufigularis – Guérin et al. 1996: 84.
Falco rufigularis – Guérin et al. 2002: 136.
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least one adult individual is present in the material. This is one of the taxa the authors highlighted as no longer living in the region today.
573. Falco cf. femoralis Temminck
Quaternary – MG
Falco femoralis (fere certe) – Winge 1887: 35.
Falconines [?] [in part] – Goeldi 1894: 76.
Falco femoralis – Lambrecht 1933: 750.
Falco femoralis – Brodkorb 1964: 294.
Falco femoralis – Mones 1986: 85.
Falco femoralis – Cuello 1988: 40.
Falco cf. femoralis – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported the remains of several individuals from Lapa da Escrivânia V, adult and young, represented by the coracoid, humerus, ulna, femur, and tarsometatarsus. The bones closely match those of Falco rufigularis but are larger and present weaker wings. They almost certainly belong to Falco femoralis . Only the distal part of the humerus, ulna, and the tarsometatarsus were compared with recent bones of a large female F. femoralis , to whom they are close in shape, although smaller. There is a marked variation in size among the bones found in the caves, but this conforms with the sex size difference of F. femoralis . Winge noted that even the smaller bones are slightly larger than the equivalent in a female F. rufigularis and that Falco deiroleucus , which does not live in the region, was not compared.
Psittaciformes Wagler
Psittacidae Rafinesque
Arinae Gray
574. Arinae indet. 1
Wiener ( 1876 ), in a text dated 1875, mentioned the complete skull of a parrot found beside a human skull fragment and complete remains of crabs in a sambaqui in the Tavares River in Ilha de Santa Catarina, Florianópolis, Santa Catarina (Casagrande 2016 ). The finding occurred during the 1875–1876 surveys that Wiener conducted by invitation of MN while he was in an expedition organized by the French government’s Ministère de l’Instruction Publique.
575. Arinae indet. 2
Quaternary – MG
Ara sp. e minoribus, præcedente [ Ara maracana et aff.] major – Winge 1887: 41.
“duas especies de Sittace ” [in part] – Goeldi 1894: 129.
Ara sp. (“kleinere und größere Formen”) [in part] – Lambrecht 1933: 770.
“undetermined macaws” [in part] – Boles 2017: 10.
Arini indet. 1 – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported the distal end of a humerus from Lapa da Escrivânia V, larger than in Primolius maracana , but smaller than in the specimen he determined as “ Ara sp. e majoribus” (see Ara sp.). He noted morphological differences between this humerus and that of a recent P. maracana , but also similarities between it and the two smaller humeri determined as fossil P. maracana . He further added that the mentioned forms are well distinguished from Cyanoliseus patagonus .
576. Arinae indet. 3
Quaternary – MG
Ara chloroptera (et forte A. hyacinthina ) [in part] – Winge 1887: 42.
[ Sittace ] hyacinthina (?) – Goeldi 1894: 129.
Ara chloroptera [in part] – Lambrecht 1933: 770.
Ara chloroptera [in part] – Brodkorb 1971: 212.
Ara chloroptera [in part] – Mones 1986: 94.
Ara chloroptera [in part] – Cuello 1988: 53.
Ara chloropterus [in part] – Boles 2017: 10.
Arini indet. 2 – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported some bones from Lapa da Lagoa do Sumidouro that are similar in size to the remains he determined as Ara chloropterus , but likely represent a species other than Ara ararauna , possibly Anodorhynchus hyacinthinus , of which he had no bones available for comparison. The material is represented by the coracoid, ulna, and femur.
577. Arinae indet. 4
Late Holocene – MG
“papagaio” – Machado et al. 1981 –1982: 209.
Machado et al. ( 1981 –1982) reported parrot bones associated with an adult woman’s skeleton (grave N° 12) from Gruta do Gentio II in Unaí, Minas Gerais.
578. Arinae indet.? 5 (spp.?)
Souza Cunha and Magalhães ( 1986 ) noted that psittacids may figure among the avian remains from Cerca Grande in Matozinhos, Minas Gerais.
579. Arinae indet. 6
Early Holocene – PE
Psittacidae – Lima 1991: 58, 6.
Lima ( 1991 ) depicted, among other remains, a psittacid beak from a burnt layer aged about 9,000 years BP at Furna do Estrago in Brejo da Madre de Deus, Pernambuco.
580. Arinae indet. 7
Schmitz et al. ( 1989 ) mentioned psittacid beaks from the GO-JA-01 rock shelter in Serranópolis, Goiás. Paulo ( 2009 ) reported that other fragments were also found.
581. Arinae indet. 8
Late Pleistocene – BA
“parrot beak” – Bryan and Gruhn 1993: 13.
Bryan and Gruhn ( 1993 ) reported a well-preserved parrot beak from the yellow silt zone near the front of the Toca dos Búzios (BA-CE-01) site in Central, Bahia. The specimen is deposited in the MN collection.
582. Arinae indet. 9
Rosa ( 2009 ) reported a bone from square A6 from the third occupation of the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. The material is deposited in the CEPA collection.
583. Arinae indet. 10 (spp.?)
Holocene – MT
Psittacidae – Pacheco 2008: 229.
Pacheco ( 2008 ) reported a complete skull and two humeri representing two small psittacids from the Santa Elina rock shelter in Jangada, Mato Grosso.
584. Arinae indet. 11
Holocene – RS
Psitacidae [sic] – Jacobus and Rosa 2013: 247.
Jacobus and Rosa ( 2013 ) reported psittacid remains from the Schneider (RS-C-14) rock shelter in São Sebastião do Caí, Rio Grande do Sul.
585. Arinae indet. 12
Holocene – RS
Psitacidae [sic] – Jacobus and Rosa 2013: 249.
Jacobus and Rosa ( 2013 ) reported psittacid remains from the Pilger (RS-C-61) rock shelter in Harmonia, Rio Grande do Sul.
586. Arinae indet. 13
Silva ( 2013 , 2014 ) reported psittacid material among remains collected by the staff of NEA-UFPE, starting in the 1990s, in rock shelters of the Pedra do Alexandre site in Carnaúba dos Dantas, Rio Grande do Norte.
Brotogeris Vigors
587. Brotogeris chiriri (Vieillot)
Quaternary – MG
Brotogerys xanthoptera – Winge 1887: 40.
Brotogerys xanthoptera – Goeldi 1894: 129.
Brotogerys xanthoptera – Lambrecht 1933: 770.
Forpus coelestis – Brodkorb 1971: 213.
Forpus coelestis [ Brotogerys xanthoptera ] – Mones 1986: 95.
Forpus coelestis – Cuello 1988: 53.
Brotogeris chiriri – Boles 2017: 10.
Brotogeris chiriri – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported several individuals from Lapa da Escrivânia V, represented by the coracoid, humerus, ulna, and femur (the last one the most uncertain). There is also a humerus from Lapa da Lagoa do Sumidouro and several bones of recent age [28] linked by soft tissues, including a humerus still attached to the scapula and the coracoid.
Brodkorb ( 1971 ) associated this record with Forpus coelestis . Regarding this, Cuello ( 1988: 53) noted that, since this species was never recorded in Brazil, “it is likely that Winge’s designation may be referred to some form of Forpus xanthopterygius ”.
The names Brotogerys xanthoptera (Spix) of Winge and the one that guided him, Conurus xanthopterus (Spix) of Reinhardt ( 1870 ) (originally Aratinga xanthopterus Spix 1824: 31, not Psittaculus xanthopterygius Spix 1824: 38) are synonymous with Brotogeris chiriri (Hellmayr 1906 ). Krabbe ( 2007 ), while reviewing the skins collected by Lund and Reinhardt, associated the name Conurus xanthopterus of Reinhardt with Brotogeris chiriri chiriri , while Forpus xanthopterygius was associated with Psittacula passerina (L.) (= Forpus passerinus ) of Reinhardt ( 1870 ).
Amazona Lesson
588. cf. Amazona sp.
Quaternary – MG
“un Perroquet ( Psittacus )” [?; in part] – Gervais 1844a: 294.
Psittacus [?; in part] – Giebel 1846: 313.
Psittacinae [?; in part] – Giebel 1847: 20.
Psittacus [?; in part] – Pictet 1853: 414.
“une espèce du groupe des perroquets” [?; in part] – Liais 1872: 303.
Psittacus aff. guyanensi [in part] – Lund (in Winge 1887 ): 40.
Chrysotis sp. e minor., vel g. aff. – Winge 1887: 40.
“duas especies de Chrysotis ” [in part] – Goeldi 1894: 129.
Chrysotis sp. (“mehrere Arten”) [in part] – Lambrecht 1933: 770.
Psittacus sp. [?; in part] – Lambrecht 1933: 770.
Chrysotis sp. [in part] – Mones 1986: 95.
“undetermined amazons” [in part] – Boles 2017: 10.
cf. Amazona sp. – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported several remains associated with a small form of this genus. From Lapa da Escrivânia V, a humerus and a tarsometatarsus that match well the record below ( Amazona sp.) but are considerably smaller. There is also an ulna, too large for this humerus, but too small for the smallest humerus of the record below. From Lapa da Escrivânia XI, material represented by the coracoid, humerus, and carpometacarpus, which matches the ones from Lapa da Escrivânia V. From a “saltpeter cave near Escrivânia”, a humerus. Finally, from an unknown locality, the distal end of a humerus (“ Psittacus aff. guyanensi ” in Lund’s catalog), very similar to the first mentioned humerus.
589. Amazona sp. 1
Quaternary – MG
Chrysotis sp., magnitudine Ch. amazonicæ etc. – Winge 1887: 40.
“duas especies de Chrysotis ” [in part] – Goeldi 1894: 129.
Chrysotis sp. (“mehrere Arten”) [in part] – Lambrecht 1933: 770.
Amazona amazonica – Brodkorb 1971: 213.
Amazona amazonica [ Chrysotis sp.” [in part]] – Mones 1986: 94–95.
Amazona amazonica – Cuello 1988: 53.
“undetermined amazons” [in part] – Boles 2017: 10.
Amazona sp. – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported some bones that closely resemble Amazona amazonica in size. From Lapa da Escrivânia V, several individuals with varying sizes are represented by the coracoid, humerus, ulna, carpometacarpus, femur, tibiotarsus, and tarsometatarsus. From Lapa da Escrivânia XI, a coracoid and an ulna.
590. Amazona sp. 2
Holocene – PR
Amazona sp. – Mendes and Rodrigues 2024: 8.
Mendes and Rodrigues ( 2024 ) associated with this genus two bones of at least one individual from Sambaqui do Toral 51 in Paranaguá, Paraná.
591. Amazona cf. aestiva (Linnaeus)
Guérin et al. ( 1996 ) reported material comparable to this species from Toca da Janela da Barra do Antonião. At least one adult and one young individual are present in the material.
Pyrrhura Bonaparte
592. Pyrrhura frontalis (Vieillot)
Quaternary – MG
Pyrrhura vittata – Winge 1887: 41.
Pyrrhura vittata – Goeldi 1894: 128.
Pyrrhura vittata – Lambrecht 1933: 770.
Pyrrhura frontalis – Brodkorb 1971: 213.
Pyrrhura frontalis [ Pyrrhura vittata ] – Mones 1986: 95.
Pyrrhura frontalis – Cuello 1988: 52.
Pyrrhura frontalis – Boles 2017: 10.
Pyrrhura frontalis – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported four humeri and two carpometacarpi from Lapa da Escrivânia V.
Anodorhynchus Spix
593. Anodorhynchus sp.
Early Holocene – MG
Psittacidae – Souza Cunha and Guimarães 1978: 219, 15.
Psittacidae – Souza Cunha and Guimarães 1981 –1982: 248, 15.
“extinct undescribed Anodorhynchus ” – Alvarenga (in Yamashita 1997 ): 181.
Anodorhynchus glaucus ou A. leari – Alvarenga 2007 b: 427–432, figs 2–3.
Anodorhynchus sp. – Nascimento and Silveira 2020: 495.
Alvarenga ( 2007 b) determined as Anodorhynchus glaucus or Anodorhynchus leari an almost complete maxilla (MNRJ-A-LV-81) from Lapa Vermelha IV in Confins, Minas Gerais, in sediments dated about 9,000 years BP. It was discovered by the Franco-Brazilian Archeological Mission (1971–1976) and briefly mentioned and depicted by Souza Cunha and Guimarães ( 1978 , 1981 –1982). A tarsometatarsus of a large bird and mammalian remains (especially rodents and marsupials) were found in the same sediments.
The identical size and shape of the specimen compared with A. glaucus or A. leari and the fact it was found between the historically known range of these two species prevent a more asserting determination. However, Alvarenga found it is quite unlikely it represents a still-unknown species, despite an earlier mention as such in Yamashita ( 1997 ). Based on the data obtained from the remains found in Bahia (see below) and Minas Gerais, and by comparing the skins and skeletons of these two species, Alvarenga reinforced the interpretation that they represent geographic races or subspecies, suggesting a contiguous geographic distribution and clinal variation of the plumage dating at least 10,000 years.
594. Anodorhynchus leari Bonaparte
Late Pleistocene – BA
Anodorhynchus leari – Alvarenga 2007 b: 427–432, figs 4–7.
In addition to the material from Minas Gerais, Alvarenga ( 2007 b) associated with this species (based on its geographical origin) bones collected by the Cástor Cartelle at Gruta dos Brejões in Morro do Chapéu, Bahia, dated about 12,200 years BP.
The material consists of an incomplete mandible (MCL A 821) and complete and well-preserved post-cranial bones, which possibly belong to a single young individual: a right coracoid (MCL A 1693), a left coracoid (MCL A 1709), a left tibiotarsus (MCL A 1707), and a right tarsometatarsus (MCL A 1769).
Eupsittula Bonaparte
595. Eupsittula aurea (Gmelin)
Quaternary – MG
Conurus aureus – Winge 1887: 41.
Conurus aureus – Goeldi 1894: 128.
Conurus aureus – Lambrecht 1933: 770.
Aratinga aurea – Brodkorb 1971: 213.
Aratinga aurea [ Conurus aureus ] – Mones 1986: 95.
Aratinga aurea – Cuello 1988: 53.
Eupsittula aurea – Boles 2017: 10.
Eupsittula aurea – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, represented by the coracoid (rather dubious), humerus, ulna, carpometacarpus, tibiotarsus, and tarsometatarsus.
This material also had its provenance erroneously listed as Lapa da Escrivânia XI by Lambrecht ( 1933 ) and as “Salpeterhule” by Brodkorb ( 1971 ).
596. Eupsittula cf. cactorum (Kuhl)
Guérin et al. ( 1996 ) reported material comparable to this species from Toca da Janela da Barra do Antonião. At least two adult individuals are present in the material.
Primolius Bonaparte
597. Primolius maracana (Vieillot)
Quaternary – MG
Ara maracana et aff. – Winge 1887: 41.
Sittace maracanã [sic] – Goeldi 1894: 129.
Ara maracana – Lambrecht 1933: 770.
Ara maracana – Brodkorb 1971: 212.
Ara maracana – Mones 1986: 94.
Ara maracana – Cuello 1988: 53.
Primolius maracana – Boles 2017: 10.
Primolius maracana – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported the distal end of a humerus and an ulna from Lapa da Escrivânia V, which differs only slightly from a recent specimen, and an incomplete humerus from Lapa da Escrivânia XI. Additionally, two other humeri that possibly came from Lapa da Escrivânia V may be attributed to this species.
Ara Lacépède
598. Ara sp.
Winge ( 1887 ) reported a humerus from Lapa da Escrivânia V, shorter than in the compared specimens of Ara ararauna , Ara macao , and Ara chloropterus , but much larger than in Primolius maracana . Additionally, the distal end of a femur from “various caves” [26] matches the humerus’ size and is about the same size as the femur of A. ararauna .
599. Ara chloropterus Gray
Quaternary – MG, RJ, PI
Ara chloroptera (et forte A. hyacinthina ) [in part] – Winge 1887: 42.
[ Sittace ] chloroptera – Goeldi 1894: 129.
Ara chloroptera [in part] – Lambrecht 1933: 770.
Ara chloroptera [in part] – Brodkorb 1971: 212.
Ara chloroptera [in part] – Mones 1986: 94.
Ara chloroptera [in part] – Cuello 1988: 53.
Ara chloropterus [in part] – Boles 2017: 10.
Ara chloropterus – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported bones of two individuals from Lapa da Escrivânia V, represented by the scapula, humerus, ulna, carpometacarpus, first phalanx of the major digit of the wing, femur, and tarsometatarsus. This material is slightly smaller than in the compared specimen of Ara chloropterus , somewhat larger than two of Ara macao , and considerably divergent in size and morphology from Ara ararauna . Additionally, there are bones of at least three individuals from Lapa da Escrivânia XI, represented by the coracoid, humerus, ulna, radius, carpometacarpus, first phalanx of the major digit of the wing, pelvis, femur, tibiotarsus, and tarsometatarsus, all slightly larger than the material of Lapa da Escrivânia V and about the same size as in A. chloropterus .
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. It is represented by very numerous remains of at least four adults and two young individuals, which suggests it was selectively hunted by the ancient inhabitants of the site. It must also have been rather abundant in the area in past times, while today it is restricted to a very small population.
“a macaw (probably A. chloroptera )” – Alvarenga (in Sick 1993 ): 261.
Ara chloroptera – Kneip et al. 1994: 51.
Psittacidae (“arara-vermelha”) – Kneip et al. 1995: 7.
“uma arara (provavelmente A. chloroptera )” – Alvarenga (in Sick 1997 ): 362.
Psittacidae (“arara-vermelha”) – Kneip et al. 1997: 19.
Psittacidae (“arara-vermelha”) – Magalhães et al. 2001: 59.
Kneip et al. ( 1994 ) reported a bone from layer I (3,610±١90 years BP) and another from layer II (3,960±200 years BP) of Sambaqui do Moa in Saquarema, Rio de Janeiro. This species no longer occurs in the region today.
Psittacara Vigors
600. Psittacara leucophthalmus (Statius Müller)
Quaternary – MG, PI
Conurus pavua – Winge 1887: 41.
[ Conurus ] pavua – Goeldi 1894: 128.
Conurus pavua – Lambrecht 1933: 770.
Aratinga leucophthalmus – Brodkorb 1971: 212.
Aratinga leucophthalmus [ Conurus pavua ] – Mones 1986: 95.
Aratinga leucophthalmus – Cuello 1988: 52.
Psittacara leucophthalmus – Boles 2017: 10.
Psittacara leucophthalmus – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported two humeri and an ulna from Lapa da Escrivânia V, a humerus from Lapa da Escrivânia XI, a carpometacarpus from a “saltpeter cave near Escrivânia”, a humerus from “various caves” [26] , and a maxilla, a humerus, and a tibiotarsus of recent age [28] .
Guérin et al. ( 1996 ) reported this species from Toca da Janela da Barra do Antonião. At least three adults and one young individual are present in the material.
Passeriformes Linnaeus
601. cf. Passeriformes 1
Holocene – PE
“aves, possivelmente passeriformes” – Silva 2003: 290.
Silva ( 2003 ) mentioned possible passeriform long bones from the Letreiro do Sobrado rock shelter in Petrolândia, Pernambuco.
602. cf. Passeriformes 2
Late Holocene – RS
cf. Passeriformes – Rosa 2006b: 232.
Rosa ( 2006b ) reported a bone from level 3 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
603. Passeriformes indet. 1 (spp.)
Quaternary – MG
“en hel Del Spurvefugle, saa godt som alle mesomyode; af de acromyode, de egenlige Sangfugle, der ellers ere de talrigste i Hulerne (smaa Finker o. s. v.), findes der netop Spor” [Lapa do Capão Seco] – Winge 1887: 11.
“Ikke faa ubestemte Spurvefugle, mesomyode og acromyode” [Lapa da Escrivânia XI] – Winge 1887: 13.
“Adskillige ubestemte Spurvefugle, mesomyode og acromyode” [Lapa da Lagoa do Sumidouro] – Winge 1887: 13.
“Nogle faa Spurvefugle, mesomyode og acromyode” [Lapa da Cerca Grande] – Winge 1887: 14.
“Enkelte Knogler af acromyode Spurvefugle” [Lapa da Escrivânia III] – Winge 1887: 14.
“Nogle faa Spurvefugle, mesomyode og acromyode” [Lapa da Escrivânia IX] – Winge 1887: 14.
“En Del ubestemte Spurvefugle, mesomyode og acromyode” [“Saltpeter cave near Escrivânia”] – Winge 1887: 14.
“Flere Spurvefugle; acromyode, i alt Fald de fleste” [Lapa do Marinho II] – Winge 1887: 14.
“En Del Spurvefugle, mesomyode og acromyode” [Lapa da Pedra dos Índios] – Winge 1887: 14.
“To Knogler af to Arter Spurvefugle” [Lapa do Periperi I] – Winge 1887: 14.
“En Del ubestemte Spurvefugle” [“various caves”] – Winge 1887: 15.
“ Passeres [”Arternes store Tal”] [in part?] – Winge 1887: 47.
Passeriformes indet. 1 (spp.) – Nascimento and Silveira 2020: 495.
Winge ( 1887 ) reported multiple indeterminate passeriforms, most from Lapa da Escrivânia V, but also “Mesomyodes” and “Acromyodes” [40] from Lapa da Escrivânia IX and XI, Lapa da Lagoa do Sumidouro, Lapa da Cerca Grande, Lapa da Pedra dos Índios, Lapa do Capão Seco (mostly “Mesomyodes”), and a “saltpeter cave near Escrivânia”, “Acromyodes” from Lapa da Escrivânia III and Lapa do Marinho II (at least the majority), two bones of two species from Lapa do Periperi I, and an amount from “various caves” [26] .
He noted that the large number of species and the uniformity of most bones make it difficult to determine them and even a much larger collection of recent, well-determined skeletons than what was available would not allow a better diagnosis of many of these specimens. Furthermore, this material may contain representatives of groups that have not been determined among the fossils collected by Lund, such as Pipridae, Parulidae, and Coerebinae.
604. Passeriformes indet. 2 (spp.)
Winge ( 1887 ) reported indeterminate “Mesomyodes” and “Acromyodes” passeriforms of recent age in a considerable number, without attribution to a locality.
605. Passeriformes indet. 3 (spp.?)
Holocene – RJ
“passeriformes (Sabiá?)” – Kneip et al. 1975: 105.
“passeriformes” – Kneip 1977: 51–52.
“passeriformes (sabia?)” – Souza Cunha et al. 1977: 147.
Passeriformes – Ruschi 1979: 36.
“aves passeriformes” – Mendonça de Souza and Mendonça de Souza 1981 –1982: 115.
Passeriformes [in part?] – Gaspar 2003: 58.
Kneip et al. ( 1975 ) reported passeriform remains (suggesting a possible association with turdids) from layers I and II of Sambaqui do Forte, in Cabo Frio, Rio de Janeiro. The bones are fragmented and consist of a few individuals.
606. Passeriformes indet. 4 (spp.)
Souza Cunha and Guimarães ( 1978 , 1981 –1982) reported numerous passeriform bones discovered by the Franco-Brazilian Archeological Mission (1971–1976) in Lapa Vermelha in Confins, Minas Gerais. The material includes mandibles, maxillae, and several long bones and is deposited in the MN collection. Their poor state of conservation (the bones fell apart easily) prevented a more accurate determination.
607. Passeriformes indet. 5 (spp.?)
Late Holocene – RJ
“passeriformes” – Kneip et al. 1984: 347.
Kneip et al. ( 1984 ) reported three fragments of small passeriforms from Sambaqui da Embratel in Rio de Janeiro, Rio de Janeiro. They figure among material collected in 1980 and 1981.
608. Passeriformes indet. 6
Late Holocene – RJ
“Fringillidae ou Thraupidae” – Lima and Silva 1984: 30.
Lima and Silva ( 1984 ) reported a complete right humerus possibly referable to Fringillidae or Thraupidae from the Ilha de Santana site in Macaé, Rio de Janeiro, dated 1,260±330 years BP. They noted that the size and morphology match that of Saltator .
609. Passeriformes indet.? 7 (spp.?)
Souza Cunha and Magalhães ( 1986 ) noted that passeriforms may figure among the avian remains from Cerca Grande in Matozinhos, Minas Gerais.
610. Passeriformes indet. 8 (spp.)
Guérin et al. ( 1993a , 1993b , 1996 ) reported indeterminate small to medium-sized passeriforms from Toca da Janela da Barra do Antonião.
611. Passeriformes indet. 9
Lima and Viana ( 1993 ) reported passeriform remains from tanks of the Lagoa da Pedra site in Conceição das Crioulas, Salgueiro, Pernambuco. Viana and Agostinho ( 1995 ) reported the remains to consist of limb parts found in the site’s facies B.
612. Passeriformes indet. 10
Late Holocene – MS
Passeriformes – Schmitz et al. 1998: 181.
Schmitz et al. ( 1998 ) reported a fragment from level 3 of the MS-CP-22 site in Ladário, Mato Grosso do Sul.
613. Passeriformes indet. 11 (spp.?)
Late Holocene – MS
Passeriformes – Schmitz et al. 1998: 187.
Schmitz et al. ( 1998 ) reported a fragment from level 2 and another from level 3 of the MS-CP-32 site in Corumbá, Mato Grosso do Sul.
614. Passeriformes indet. 12 (spp.?)
Dias ( 2003 , 2004a ) reported small passeriform remains from the Sangão (RS-S-327) rock shelter in Santo Antônio da Patrulha, Rio Grande do Sul.
615. Passeriformes indet. 13 (spp.?)
Holocene – RS
Passeriformes – Jacobus 2004: 99.
Jacobus ( 2004 ) reported passeriform remains from the Dalpiaz (RS-LN-1) rock shelter in Maquiné, Rio Grande do Sul. The material is deposited in the MARSUL collection.
616. Passeriformes indet. 14
IBAMA ( 2005b ) reported passeriform remains from Lapa do Carlúcio in Itacarambi, Minas Gerais. Oliveira ( 2008 ) reported an almost complete subfossil skeleton from the same cave, but it is unclear whether it refers to the same material.
617. Passeriformes indet. 15
Late Holocene – RS
Emberizidae – Rosa 2006c: 250.
Rosa ( 2006c ) reported an “Emberizidae” bone from level 1 of square 3 of the RS-LC-80 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
618. Passeriformes indet. 16 (spp.?)
Rosa ( 2009 ) reported 179 bones (four from square A6, one from B6, one from B7, 50 from C5, three from C6, 29 from D5, 88 from D6, and three from D7) from the Garivaldino (RS-TQ-58) site in Montenegro, Rio Grande do Sul. Remains from square A6 were detected in the site’s three occupation periods. The author mentioned “Emberizidae” among the determined material. The material is deposited in the CEPA collection.
619. Passeriformes indet. 17
Teixeira ( 2006 ) reported a bone from level 7 of the Rincão (SC-IÇ-06) site in Içara, Santa Catarina. The material is deposited in the IAP/Unisinos collection.
620. Passeriformes indet. 18
Late Holocene – RS
“Passeriformes não-identificado” – Rosa et al. 2009: 142.
Rosa et al. ( 2009 ) reported two bones of at least one individual from the Guarani site RS-T-114 in Marques de Souza, Rio Grande do Sul.
621. Passeriformes indet. 19
Late Holocene – MS
Passeriformes – Schmitz et al. 2009: 331.
Schmitz et al. ( 2009 ) reported a bone from level 7 of the MS-MA-180 site in Corumbá, Mato Grosso do Sul.
622. Passeriformes indet. 20
Late Holocene – MS
Passeriformes – Schmitz et al. 2009: 334.
Schmitz et al. ( 2009 ) reported a bone from level 6 of the MS-MA-84 site in Corumbá, Mato Grosso do Sul.
623. Passeriformes indet. 21
Late Holocene – MS
Passeriformes – Schmitz et al. 2009: 342.
Schmitz et al. ( 2009 ) reported a bone from level 3 of the MS-MA-16A site in Corumbá, Mato Grosso do Sul.
624. Passeriformes indet. 22 (3 spp.)
Quaternary – BA
“três espécies ainda não determinadas [Passeriformes]” – Silva and Cozzuol 2010: 111.
Silva and Cozzuol ( 2010 ) reported eight humeri representing three indeterminate passeriforms among material collected in the 1980s at Toca da Boa Vista in Campo Formoso, Bahia. The fossils are deposited in the MHNJB/UFMG collection.
625. Passeriformes indet. 23
Silva ( 2013 , 2014 ) reported passeriform material among remains collected by the staff of NEA-UFPE, starting in the 1990s, in rock shelters of the Pedra do Alexandre site in Carnaúba dos Dantas, Rio Grande do Norte.
626. Passeriformes indet. 24
Late Holocene – RS
Passeriformes – Ferrasso and Schmitz 2015: 126.
Ferrasso and Schmitz ( 2015 ) reported a passeriform phalanx about 7 mm long from the RS-LN-279 site at the northern coast of Rio Grande do Sul, aged 3,310±40 years BP.
Thamnophilidae Swainson
Thamnophilinae Swainson
Dysithamnus Cabanis
627. Dysithamnus sp.
Quaternary – TO
Desithamnus [sic] sp. – Salum et al. 2016: 153.
Salum et al. ( 2016 ) reported incomplete post-cranial thamnophilid materials from Gruta do Urso ( Fig. 1 .17) in Aurora do Tocantins, Tocantins, aged between 22,000–3,800 years BP. The remains are represented by the humeri (the most abundant, with four specimens), tibiotarsi, tarsometatarsi, and femora. Five genera were preliminarily determined, but the remains likely include a greater diversity. One of the determined genera is Dysithamnus .
Herpsilochmus Cabanis
628. Herpsilochmus sp.
Quaternary – TO
Herpsilochmus sp. – Salum et al. 2016: 153.
Salum et al. ( 2016 ) reported this genus among post-cranial remains from Gruta do Urso, Aurora do Tocantins, Tocantins.
Thamnophilus Vieillot
629. Thamnophilus sp.
Quaternary – TO
Thamnophilus sp. – Salum et al. 2016: 153.
Salum et al. ( 2016 ) reported this genus among post-cranial remains from Gruta do Urso, Aurora do Tocantins, Tocantins.
Pyriglena Cabanis
630. Pyriglena sp.
Quaternary – TO
Pyriglena sp. – Salum et al. 2016: 153.
Salum et al. ( 2016 ) reported this genus among post-cranial remains from Gruta do Urso, Aurora do Tocantins, Tocantins.
Hypocnemis Cabanis
631. Hypocnemis sp.
Quaternary – TO
Hypocnemis sp. – Salum et al. 2016: 153.
Salum et al. ( 2016 ) reported this genus among post-cranial remains from Gruta do Urso, Aurora do Tocantins, Tocantins.
Formicariidae Gray
632. Formicariidae indet. (spp.)
Winge ( 1887 ) reported remains of multiple species, but it was not possible to safely determine them beyond the family level. Few recent skeletons were available for comparison and no exact matches were found for the humeri of Pyriglena leucoptera and Formicivora rufa rufatra —thamnophilids then included in Formicariidae (which indicates that the material under this determination may include more than one family). Two small humeri from Lapa da Escrivânia V are especially thin and delicate even for this family; they have the same length as observed in Troglodytes musculus , but the humerus of this species is relatively more robust.
Goeldi ( 1894 ) erroneously listed Pyriglena leucoptera as provisionally represented among the material.
Chamaeza Vigors
633. Chamaeza campanisona (Lichtenstein)
Quaternary – MG
Chamæzosa brevicauda (et aff.) – Winge 1887: 47.
Chamaezasa brevicauda – Goeldi 1894: 363.
Chamaezosa brevicauda – Lambrecht 1933: 775.
Chamaeza – Storer 1960: 81.
Chamaeza campanisona – Fisher 1967: 753.
Chamaeza brevicauda – Brodkorb 1978: 147.
Chamaeza brevicauda – Mones 1986: 98.
Chamaeza brevicauda – Cuello 1988: 62.
Chamaeza brevicauda – Unwin 1993: 731.
Chamaeza campanisona – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported a coracoid from Lapa da Escrivânia XI and seven humeri from Lapa da Escrivânia V, five of which match the ones of recent specimens and two slightly divergent and smaller. He noted that this species’ morphology is easily distinguishable from the other “Mesomyodes” passeriforms, but, from the “ Grallaria group” [41] , it was the only one with available recent bones.
Dendrocolaptidae Gray
634. Dendrocolaptidae indet. (spp.)
Quaternary – MG
Anabates [?; in part] – Lund 1841d: 18.
Dendrocalaptes [sic] [?; in part] – Lund 1841d: 18.
Opetiorrhynchus [sic] [?; in part] – Lund 1841d: 18.
Anabates poliocephalus [?; in part] – Lund 1841d: 18.
Dendrocalaptes [sic] [?; in part] – Giebel 1846: 312.
Anabates ? [?; in part] – Giebel 1846: 313.
Anabates [?; in part] – Giebel 1847: 13.
Opetiorhynchus [?; in part] – Giebel 1847: 13.
Dendrocalaptes [sic] [?; in part] – Giebel 1847: 17.
Anabates [?; in part] – Pictet 1853: 412.
Opetiorhynchus [?; in part] – Pictet 1853: 412.
Dendrocolaptes [?; in part] – Pictet 1853: 414.
“des Grimpars ( Anabates de Temminck)” [?; in part] – Liais 1872: 303.
“les Grimpereaux” [?; in part] – Liais 1872: 303.
Furnarius [?; in part] – Reinhardt 1881: 142.
Dendrocolaptidæ [“Species multæ, indeterminatæ, etiam affines Furnario etc.”] – Winge 1887: 8, 47–48.
Dendrocolaptides [in part] – Goeldi 1894: 363.
Anabates [?; in part] – Lambrecht 1933: 779.
Opetirhynchus [sic] [?; in part] – Lambrecht 1933: 779.
“2 indet. spp. of different genera” [in part] – Fisher 1967: 753.
Dendrocolaptidae indet. (spp.) – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported multiple indeterminate species. He noted that few recent skeletons were available for comparison, and none of the most similar humeri found in the caves is a good match to Lochmias nematura , Synallaxis cinnamomea , Sittasomus griseicapillus sylviellus , and Dendroma rufa . Many are similar to Furnarius rufus albogularis but a little more robust in length when compared with a recent specimen.
Brodkorb ( 1978 ) associated with this indeterminate material the names Anabates (= Synallaxis ), Dendrocalaptes [sic], and Opetiorrhynchus [sic] ( Opetiorhynchus ; = Furnarius ) used by Lund ( 1841d ).
Dendrocolaptinae Gray
Xiphocolaptes Lesson
635. Xiphocolaptes spp.?
Quaternary – BR
“ Xiphocolaptes species” – Alvarenga (in Marantz et al. 2003 ): 358.
In addition to listing the records reported by Winge ( 1887 ) from the Lagoa Santa region, Marantz et al. ( 2003 ), through information received from Herculano Alvarenga, noted that “specimens of Xiphocolaptes species have been found at another site in Brazil”, without further details. Perhaps this is the same record reported by Sales ( 2003 ) (see below).
636. Xiphocolaptes falcirostris (Spix)
Quaternary – MG
Xiphocolaptes franciscanus sp. – Sales 2003: 214.
Sales ( 2003 ) reported twelve specimens among the bones and mummified carcasses from Zone II of Lapa do Rezar in Itacarambi, Minas Gerais.
637. Xiphocolaptes albicollis (Vieillot)
Quaternary – MG
Xiphocolaptes albicollis – Winge 1887: 48.
Xiphocolaptes albicollis – Goeldi 1894: 363.
Xiphocolaptes albicollis – Lambrecht 1933: 775.
Xiphocolaptes albicollis – Fisher 1967: 753.
Xiphocolaptes albicollis – Brodkorb 1978: 146.
Xiphocolaptes albicollis – Mones 1986: 98.
Xiphocolaptes albicollis – Cuello 1988: 61.
Xiphocolaptes albicollis – Marantz et al. 2003: 358.
Xiphocolaptes albicollis – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported a tarsometatarsus from Lapa da Escrivânia V and a tibiotarsus from an unknown locality.
Lepidocolaptes Reichenbach
638. Lepidocolaptes angustirostris (Vieillot)
Quaternary – MG
Picolaptes bivittatus – Winge 1887: 48.
Picolaptes bivittatus – Goeldi 1894: 363.
Picolaptes bivittatus – Lambrecht 1933: 775.
Lepidocolaptes angustirostris – Fisher 1967: 753.
Lepidocolaptes angustirostris – Brodkorb 1978: 146.
Lepidocolaptes angustirostris [ Picolaptes bivittatus ] – Mones 1986: 98.
Lepidolaptes angustirostris [sic] – Cuello 1988: 62.
Lepidocolaptes angustirostris – Marantz et al. 2003: 358.
Lepidocolaptes angustirostris – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported a well-matching tibiotarsus from Lapa da Escrivânia V, as well as a skull and most of the other bones of an individual of recent age [28] .
Furnariidae Gray
Furnariinae Gray
Furnarius Vieillot
639. cf. Furnarius rufus (Gmelin) 1
Schmitz et al. ( 1998 ) reported three fragments comparable to this species and attributable to an individual from level 7 of the MS-CP-20 site in Corumbá, Mato Grosso do Sul.
640. cf. Furnarius rufus (Gmelin) 2
Late Holocene – RS
cf. Furnarius rufus – Silva and Rosa 2006: 220.
Silva and Rosa ( 2006 ) reported a bone comparable to this species from the RS-LC-82 site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
641. Furnarius rufus (Gmelin)
Quaternary – MG
Furnarius rufus – Sales 2003: 214.
Sales ( 2003 ) reported a specimen among the bones and mummified carcasses from Zone II of Lapa do Rezar in Itacarambi, Minas Gerais.
Tyrannidae Vigors
642. Tyrannidae indet. 1 (spp.)
Winge ( 1887 ) reported multiple species, but it was not possible to determine them beyond the family level. He noted that the shape of the bones is very similar between them.
643. Tyrannidae indet. 2
Late Holocene – RJ
Tyranídae [sic] – Gaspar 2003: 58.
Gaspar ( 2003 ) reported tyrannid material from the Ponta da Cabeça site in Arraial do Cabo, Rio de Janeiro.
Tyranninae Vigors
Pitangus Swainson
644. Pitangus sulphuratus (Linnaeus)
Rosa ( 2004 ) reported a bone from the Serranópolis phase (early–late Holocene) square 16H of the GO-JA-01 rock shelter in Serranópolis, Goiás. The material is deposited in the IAP/Unisinos collection.
Pitangus sulphuratus – Silva and Cozzuol 2010: 111.
Silva and Cozzuol ( 2010 ) reported an incomplete tarsometatarsus lacking the proximal epiphysis among material found in the 1980s at Toca da Boa Vista in Campo Formoso, Bahia. The fossil is deposited in the MHNJB/UFMG collection.
Megarynchus Thunberg
645. Megarynchus pitangua (Linnaeus)
Winge ( 1887 ) reported a skull of recent age without attributing it to a specific locality.
Myiozetetes Sclater
646. Myiozetetes similis (Spix)
Quaternary – MG
Myiozetetes similis sp. – Sales 2003: 214.
Sales ( 2003 ) reported a specimen among the bones and mummified carcasses from Zone II of Lapa do Rezar in Itacarambi, Minas Gerais.
Vireonidae Swainson
647. Vireonidae indet.
Winge ( 1887 ) reported a humerus from Lapa da Escrivânia V, significantly smaller than in the following record, and another similar humerus of recent age [28] .
Cyclarhis Swainson
648. Cyclarhis sp.
Quaternary – MG
Cyclorhis ? sp. [sic] – Winge 1887: 49.
Cyclorhis esp. – Goeldi 1894: 363.
Cyclorhis sp. (?) [sic] – Lambrecht 1933: 780.
Cyclarhis gujanensis – Fisher 1967: 758.
Cyclarhis gujanensis (?) – Brodkorb 1978: 195.
Cyclarhis – Sick 1984b: 642.
Cyclarhis gujanensis [? Cyclorhis sp.] – Mones 1986: 99–100.
Cyclarhis gujanensis – Cuello 1988: 68.
Cyclarhis – Sick 1993: 553.
Cyclarhis gujanensis – Unwin 1993: 733.
Cyclarhis – Sick 1997: 714.
Cyclarhis sp. – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported three humeri from Lapa da Escrivânia V, all very similar to but slightly smaller than Cyclarhis gujanensis . There is also a similar humerus from an unknown locality.
Corvidae Leach
Cyanocorax Boie
649. Cyanocorax cristatellus (Temminck)
Quaternary – MG
Dendrocalaptes [sic] [?; in part] – Lund 1841d: 18.
Dendrocalaptes [sic] [?; in part] – Giebel 1846: 312.
Dendrocalaptes [sic] [?; in part] – Giebel 1847: 17.
Dendrocolaptes [?; in part] – Pictet 1853: 414.
Dendrocolaptes aff. gujanensi [in part] – Lund (in Winge 1887 ): 48.
Cyanocorax cyanoleucus – Winge 1887: 48.
Cyanocorax cyanoleucus – Goeldi 1894: 363.
Cyanocorax cyanoleucus – Lambrecht 1933: 783.
Cyanocorax cristatella – Brodkorb 1978: 161.
Cyanocorax cristatella [ Cyanocorax cyanoleucus ] – Mones 1986: 99.
Cyanocorax cristatellus – Cuello 1988: 69.
Cyanocorax cristatellus – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported remains from various sites. From Lapa da Escrivânia V, the proximal end of a scapula, the distal end of a humerus, and the first phalanx of the left hallux, which broke when the bird was alive and became slightly shorter after calcifying. From a “saltpeter cave near Escrivânia”, a tarsometatarsus. From “various caves” [26] , a maxilla and the distal end of a tibiotarsus, which is well distinguished from that of Cacicus , Pyroderus , and Procnias . Finally, from a “cave near Sumidouro” (other than Lapa da Lagoa do Sumidouro), the proximal end of a humerus of a Cyanocorax with an uncertain determination (“ Dendrocolaptes aff. gujanensi ” in Lund’s catalog; perhaps the material Lund 1841d referred to as Dendrocalaptes [sic]). It is smaller than in Cyanocorax cristatellus and larger than and divergent from Cyanocorax cyanopogon , the two species Winge had available for comparison.
Hirundinidae Rafinesque
650. Hirundinidae indet. 1
Winge ( 1887 ) reported bones of several individuals from Lapa da Escrivânia V, a humerus from Lapa da Escrivânia XI, and bones from several individuals of recent age [28] . The total material is represented by the humerus, ulna, femur, and tibiotarsus, and perhaps represents two species. There are similarities with Pygochelidon cyanoleuca and Tachycineta leucorrhoa .
651. Hirundinidae indet. 2
Late Holocene – RS
Hirundinidae – Rosa 2006b: 232.
Rosa ( 2006b ) reported two bones from level 1 of the Chácara do Leão (RS-LC-96) site in Palmares do Sul, Rio Grande do Sul. The material is deposited in the IAP/Unisinos collection.
Progne Boie
652. Progne chalybea (Gmelin)
Quaternary – MG
Hirundo ( Progne ) domestica – Winge 1887: 48.
Progne domestica – Goeldi 1894: 248.
Chelidon rustica [in part?] – Lambrecht 1933: 776.
Progne chalybea – Brodkorb 1978: 154.
Progne chalybea [ Hirundo ( Progne ) domestica ] – Mones 1986: 98–99.
Progne chalibea [sic] – Cuello 1988: 63.
Progne chalybea – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, a humerus from Lapa da Escrivânia XI, and another humerus from Lapa da Lagoa do Sumidouro. The total material is represented by the humerus, ulna, carpometacarpus, first phalanx of the major digit of the wing, femur, and tarsometatarsus, with significant individual variations.
Hirundo Linnaeus
653. Hirundo sp.
Quaternary – MG
Hirundo sp. indet., Hirundine rustica paullo major – Winge 1887: 48.
“pouco maior do que a H. rustica da Europa” – Goeldi 1894: 248.
Chelidon rustica [in part?] – Lambrecht 1933: 776.
Hirundo sp. [in part?] – Lambrecht 1933: 776.
Hirundo rustica (?) – Brodkorb 1978: 155.
Hirundo rustica – Mones 1986: 98.
Hirundo rustica – Cuello 1988: 63.
Hirundo sp. – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported several bones from Lapa da Escrivânia V, represented by the humerus, ulna, and carpometacarpus, all slightly larger than and somewhat divergent from those of Hirundo rustica .
Lambrecht ( 1933 ), as pointed out by Brodkorb ( 1978 ), associated material from Lapas da Escrivânia V and XI and Lapa da Lagoa do Sumidouro to Chelidon rustica . Winge described materials from these localities together as Hirundo ( Progne ) domestica (= Progne chalybea ; see above).
Troglodytidae Swainson
654. Troglodytidae indet.
Winge ( 1887 ) reported a humerus from Lapa da Escrivânia V and another humerus of recent age [28] , of which a determination beyond the family level was not possible. He noted they are much smaller than in the following record.
Troglodytes Vieillot
655. Troglodytes musculus Naumann
Winge ( 1887 ) reported several humeri from Lapa da Escrivânia V and a single humerus of recent age [28] .
Turdidae Rafinesque
Turdus Linnaeus
656. Turdus sp. 1
Winge ( 1887 ) associated with this genus some humeri from Lapa da Escrivânia V.
657. Turdus sp. 2
Winge ( 1887 ) reported a skull of recent age without attributing it to a specific locality.
Mimidae Bonaparte
Mimus Boie
658. Mimus saturninus (Lichtenstein)
Winge ( 1887 ) reported several humeri from Lapa da Escrivânia V, a humerus from Lapa da Lagoa do Sumidouro, and two humeri of recent age [28] .
Passerellidae Cabanis & Heine
Arremon Vieillot
659. Arremon flavirostris Swainson
Winge ( 1887 ) reported a skull of recent age without attributing it to a specific locality.
Icteridae Vigors
660. Icteridae indet. (spp.)
Winge ( 1887 ) reported multiple indeterminate passeriform bones of various species from Lapa da Escrivânia V and a considerable number from other caves. He noted that, of skulls and other more diagnostic parts, only the cave materials were available, and, in general, the limb bones of Passeriformes are quite similar, with only some easily recognizable forms determined. Of the recognized families of the Brazilian avifauna by Pacheco et al. ( 2021 ), he cited Icteridae and Thraupidae.
Cacicinae Bonaparte
Psarocolius Wagler
661. Psarocolius decumanus (Pallas)
Quaternary – MG
Cassicus ( Ostinops ) cristatus – Winge 1887: 50.
Ostinops cristatus – Goeldi 1894: 363.
Cassicus ( Ostinops ) cristatus – Lambrecht 1933: 785.
Psarcolius decumanus – Brodkorb 1978: 200.
Psarocolius decumanus [ Cassicus ( Opstinops ) cristatus ] – Mones 1986: 99.
Psarocolius decumanus – Cuello 1988: 68.
Psarocolius decumanus – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported the proximal end of a scapula from a “saltpeter cave near Escrivânia”, which fits well to this species. It differs somewhat from that of Cyanocorax and is larger than that of Cyanocorax cristatellus (it has a different aspect from the tarsometatarsus of this species found in the same cave). It also differs from the scapula of large cotingids such as Pyroderus and Procnias .
Agelaiinae Swainson
Gnorimopsar Richmond
662. Gnorimopsar chopi (Vieillot)
Quaternary – MG
Aphobus chopi – Winge 1887: 50.
Aphobus chopi – Goeldi 1894: 363.
Aphobus Chopi [sic] – Lambrecht 1933: 790.
Gnorimopsar chopi – Brodkorb 1978: 201.
Gnorimopsar chopi [ Aphobus chopi ] – Mones 1986: 99.
Gnorimopsar chopi – Cuello 1988: 69.
Gnorimopsar chopi – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported a maxilla and humeri from Lapa da Escrivânia V.
Pseudoleistes Sclater
663. Pseudoleistes guirahuro (Vieillot)
Quaternary – MG
Pseudoleistes viridis – Winge 1887: 50.
Pseudoleistes viridis – Goeldi 1894: 363.
Pseudoleistes viridis – Lambrecht 1933: 790
Pseudoleistes guirahuro – Brodkorb 1978: 201.
Pseudoleistes guirahuro [ Pseudoleistes viridis ] – Mones 1986: 99.
Pseudoleistes guirahuro – Cuello 1988: 69.
Pseudoleistes guirahuro – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported humeri from Lapa da Escrivânia V.
Cardinalidae Ridgway
Piranga Vieillot
664. Piranga flava (Vieillot)
Winge ( 1887 ) reported a maxilla of recent age without attributing it to a specific locality.
Thraupidae Cabanis
665. Thraupidae indet. 1 (spp.)
Quaternary – MG
Tanagridæ (og Coerebidæ), Emberizidæ (med Pitylinæ o. s. v.) og Icteridæ [“Familiæ characteribus ossium membrorum inter se non distinctæ. Species permultæ indeterminatæ.”] [in part] – Winge 1887: 8, 49–50.
Tanagrides [in part] – Goeldi 1894: 363.
Thraupidae indet. 1 (spp.) – Nascimento and Silveira 2020: 496.
Winge ( 1887 ) reported thraupid (possibly including Coerebinae) remains among indeterminate materials from Lapa da Escrivânia V and other caves.
666. Thraupidae indet. 2
Winge ( 1887 ) reported a skull, which he described as somewhat different from that of Coereba flaveola chloropyga . The material is of recent age and was not attributed to a specific cave.
Dacninae Sundevall
Tersina Vieillot
667. Tersina viridis (Illiger)?
Quaternary – MG
Procnias tersa ? – Winge 1887: 50.
Procnias tersa – Goeldi 1894: 363.
Procnias tersa (?) – Lambrecht 1933: 775.
Tersina viridis – Storer 1960: 89.
Tersina viridis – Fisher 1967: 758.
Tersina viridis – Brodkorb 1978: 198.
Tersina viridis [ Procnias tersa ?] – Mones 1986: 99.
Tersina viridis – Cuello 1988: 67.
Tersina viridis ? – Nascimento and Silveira 2020: 497.
Winge ( 1887 ) associated, with uncertainty, the fragment of a maxilla from Lapa da Escrivânia V with this species.
Although Winge mentioned that no cotingid remains were found in the caves, Storer ( 1960 ) commented that the original description of the material under the genus Procnias caused Lambrecht ( 1933 ) and subsequent authors to erroneously list it as cotingid, arguing that this genus had been used for neotropical bellbirds since 1907. However, Lambrecht appears to have only listed the material as presented by Winge (as “ Procnias tersa Auct.”), similarly to most other records in his study.
Saltatorinae Bonaparte
Saltator Vieillot
668. Saltator similis d’Orbigny & Lafresnaye
Winge ( 1887 ) reported a maxilla from Lapa da Escrivânia V.
Thraupinae Cabanis
Thraupis Boie
669. Thraupis sayaca (Linnaeus)
Winge ( 1887 ) reported a maxilla of recent age without attributing it to a specific locality.
CONCLUSION
A complete panorama of the avian fossil and subfossil diversity in Brazil is far from being reached, but herein we presented a list with the most important records in both paleontological and archeological literature. This list is naturally biased by several biological, geological, political, and practical factors, besides our more conservative approach to the classification of the records, but, to date, it is the most thorough compilation on this subject for the country.
Our research resulted in 669 record units (not necessarily unique taxa), which can be divided into footprints (1), coprolites (21), feathers (169), eggs (4), and osteological remains (474) (Tables 5 and 6 ).
The following sections analyze this compilation from different perspectives.
Taxonomic representation
Of the 669 record units, 355 were sourced from the paleontological literature, 217 from the archeological literature, 29 from both (when sub-records are present), and 68 were firsthand accounts. Osteological materials are distributed in 77 higher taxa (families or equivalents). Multiple sub-records are present in 47 records. Uncertain taxonomical determination (“indet.”, “cf.”, “aff.”, “?”) is present in 509 records. Records that include or most likely include more than one certain taxon (“spp.”, “spp.?”) account for 38. Most records belong to living or non-diagnosable taxa (417). Several records may represent a single species, and other materials may comprise disparate taxa under a common determination.
Columbidae is the most exclusive clade with the best taxonomical representation in the fossil and subfossil records (33) and is followed by Rallidae (32) and Psittacidae (27).
The following orders of living native Brazilian avifauna (following Pacheco et al. 2021 ) do not have published fossil or subfossil representatives found during this research: Steatornithiformes, Eurypygiformes, and Phaethontiformes. In the same category, the following families do not have published fossil or subfossil representatives: Steatornithidae, Psophiidae, Heliornithidae, Haematopodidae, Recurvirostridae, Burhinidae, Chionidae, Thinocoridae, Rostratulidae, Glareolidae, Stercorariidae, Eurypygidae, Phaethontidae, Oceanitidae, Hydrobatidae, Pelecanidae, Pandionidae, Galbulidae, Capitonidae, Melanopareiidae, Conopophagidae, Grallariidae, Rhinocryptidae, Scleruridae, Xenopidae, Pipridae, Cotingidae, Tityridae, Oxyruncidae, Onychorhynchidae, Pipritidae, Platyrinchidae, Tachurisidae, Rhynchocyclidae, Polioptilidae, Donacobiidae, Motacillidae, Fringillidae, Parulidae, and Mitrospingidae.
None of the published records includes families with only living representatives outside Brazil. Of globally extinct families, the following have fossil representatives in the country: Quercymegapodiidae (2), Pelagornithidae (1; dubious), Palaelodidae (1), Teratornithidae (1), Gracilitarsidae (1), and Phorusrhacidae (3).
New described fossil taxa include 15 genera and 20 species of seven extant and 10 extinct families (or equivalent clades): Cratoavis cearensis , Kaririavis mater , Diogenornis fragilis , Chaunoides antiquus , Neochen pugil , Taubacrex granivora , Ameripodius silvasantosi , Agnopterus sicki , Hoazinavis lacustris , Ciconia lydekkeri , Anhinga minuta , Macranhinga ranzii , Brasilogyps faustoi , Wingegyps cartellei , Pleistovultur nevesi , Taubatornis campbelli , Eutreptodactylus itaboraiensis , Paleopsilopterus itaboraiensis , Itaboravis elaphrocnemoides , and Paraphysornis brasiliensis . The attribution of Pelagornis longirostris to Brazil is problematic but is included here for historical completeness. Additionally, two extinct species named from elsewhere have been found in the country, though one attribution is provisional ( Palaelodus aff. ambiguus ) and the other ( Rhea fossilis ) is possibly synonymous with a still living species. Life reconstructions of the extinct taxa with at least a generic attribution are presented in Figs 30–35, including approximate scale.
Taxonomic representation among osteological remains listed in the main text. Records with multiple, indeterminate species (“spp.”) are listed as a single unit here. BR = Brazil, for records without a defined state of origin.
Temporal representation
The 669 record units spawn from the Early Cretaceous to the latest Holocene, but several temporal gaps remain to be filled. Mesozoic remains are represented in 166 Early Cretaceous and 6 Late Cretaceous records. Cenozoic remains have 12 records in the early Eocene, 1 in the middle Eocene, 1 in the Eocene/Oligocene, 41 in the late Oligocene/early Miocene, 1 in the early Miocene, 16 in the late Miocene, and 424 in the Quaternary, ranging from the early Pleistocene to pre-colonial times, besides 1 possible Paleogene record without recorded provenance.
Early Cretaceous remains are dominated by feathers, with just a handful of osteological specimens attributed to enantiornitheans and ornithuromorphs known to exist. Late Cretaceous records hold enantiornitheans and other indeterminate remains, besides the only known Mesozoic fossil egg. Few early Paleogene records are known, and the determined taxa consist mostly of medium-sized birds of terrestrial habit. The Paleogene–Neogene transition holds most of the diversity of described extinct taxa, including very large species, of terrestrial and aquatic habit, besides eggs, coprolites, and footprints, of which the last two are the only known records of this type for the country. The known late Miocene avifauna consists mostly of anhingids. Remains from caves make up the bulk of Quaternary remains, with representatives in all its stages, from paleontological and archeological sites, and most of the known living groups are represented. The most complete fossil avifauna known for the country is the one from the Quaternary of the Lagoa Santa Karst. In sum, the known past avian record in Brazil is essentially modern, even if we exclude zooarcheological remains.
Life reconstruction of named Early Cretaceous taxa from the Crato Formation: the enantiornithean Cratoavis cearensis (A) and the ornithuromorph Kaririavis mater (B) (R.S. Nascimento). Scale bar: 10 mm.
Life reconstruction of named Early Eocene taxa from the São José de Itaboraí Basin: the ratite Diogenornis fragilis (A), the gracilitarsid Eutreptodactylus itaboraiensis (B), and the cariamiforms Itaboravis elaphrocnemoides (C) and Paleopsilopterus itaboraiensis (D) (R.S. Nascimento). Scale bar: 100 mm (A, D), 15 mm (B), 50 mm (C).
Life reconstruction of named Late Oligocene/Early Miocene taxa from the Tremembé Formation: the opisthocomid Hoazinavis lacrustris (A), the phorusrhacid Paraphysornis brasiliensis (B), the cathartid Brasilogyps faustoi (C), and the teratornithid Taubatornis campbelli (D) (R.S. Nascimento). Scale bar: 50 mm (A, C, D), 300 mm (B).
Life reconstruction of named Late Oligocene/Early Miocene taxa from the Tremembé Formation: the quercymegapodiids Taubacrex granivora (A) and Ameripodius silvasantosi (B), the phoenicopterid Agnopterus sicki (C), and the palaelodid Palaelodus aff. ambiguus (D) (R.S. Nascimento). Scale bar: 50 mm.
Life reconstruction of various Cenozoic taxa: the anhimid Chaunoides antiquus (A) from the Tremembé Formation (Late Oligocene/Early Miocene), the anhingids Anhinga minuta (B) and Macranhinga ranzii (C) from the Solimões Formation (Late Miocene), and the rallid Rallus sp. (D) from Fernando de Noronha (Late Holocene) (R.S. Nascimento). Scale bar: 50 mm (A, B, D), 100 mm (C).
Life reconstruction of named Quaternary taxa from cave deposits: the anatid Neochen pugil (A), the ciconiid Ciconia lydekkeri (B), and the cathartids Pleistovultur nevesi (C) and Wingegyps cartellei (D) (R.S. Nascimento). Scale bar: 100 mm.
Spatial representation
The political territory defined as Brazil currently has 26 states and the Federal District distributed into five Regions (North, Northeast, Central-West, Southeast, and South). Avian remains are known from all Regions.
The 669 record units (and its sub-records) were assigned to the following states: North: Acre (14), Amazonas (3), Pará (2), Rondônia (1), Tocantins (5); Northeast: Alagoas (1), Bahia (16), Ceará (171), Pernambuco (12), Piauí (38), Rio Grande do Norte (10), Sergipe (4); Central-West: Goiás (20), Mato Grosso (4), Mato Grosso do Sul (13); Southeast: Espírito Santo (1), Minas Gerais (179), Rio de Janeiro (60), São Paulo (54); South: Paraná (5), Rio Grande do Sul (85), and Santa Catarina (38). Additionally, 2 records lack a defined state of origin and 1 was dubiously attributed to the country. No catalog record was attributed to Amapá and Roraima, but indeterminate mentions are presented in Table 4 . Paraíba and the Federal District are the only territories without any mention found during our research.
Pre-Quaternary records are mostly restricted to individual outcrops. Early Cretaceous records are restricted to Ceará, and Late Cretaceous to São Paulo and Minas Gerais. Most Paleogene records come from Rio de Janeiro, with meager records from Paraná and Minas Gerais. Except for some late Miocene records of Acre and Amazonas, the late Cenozoic records are virtually restricted to remains of Quaternary age. Paleontological materials come predominantly from Minas Gerais, Bahia, and Piauí, while most archeological ones come from the Southeast and South regions, notably from Rio Grande do Sul and Rio de Janeiro. Rhea americana and its eggs are the record with the widest occurrence, recorded in eight states of all regions except North.
Final considerations
As Bittencourt and Langer ( 2012 ) pointed out concerning the few non-avian dinosaur species described for Brazil, the small number of fossil bird species known is also incongruent in the face of the country’s continental dimensions and its rich sedimentary deposits, even considering the fragile nature of avian bones. Several initiatives could help develop further paleornithological and archeornithological studies in Brazil, including, as for science in general, better investments leading to new fieldwork campaigns as well as the creation of a more stimulating scenario for new researchers, reanalysis of museum collections (in which avian bones, generally poor preserved, go unnoticed) and expansion of reference collections, law enforcement on fossil traffic (which led valuable species into the scientific limbo of private collections), and preservation of geological sites.
The compilation and update on the knowledge of Brazilian fossil and subfossil provide a basis for further studies, whether at a regional or national scale. Apart from expanding the understanding of avian evolution and taxonomic diversity, this knowledge is an essential source of paleobiogeographic and paleoenvironmental information (Alvarenga and Höfling 2000 ) and may also help in future strategies for managing living species (Rosa 2008b , Dietl and Flessa 2011 ).
ACKNOWLEDGMENTS
The intention to cover most of the bird fossil and subfossil records known for Brazil would never have been possible without the help of numerous people. We first thank Piter Kehoma Boll and Carlos Augusto Chamarelli for their tireless help with translations, reviews, suggestions, acquisition of bibliography, and inspiring discussions. Various professionals took part of their time to help us with requests for information, photographs, and bibliographic references or received us in the collection for which they were responsible. We thank Kasper Lykke Hansen (ZMUC), Sandra Chapman (NHMUK), Fred Ruhe (BirdForum), Gustavo M.E.M. Prado (IGc/USP), Glauco Freire de Albuquerque (UFRN), Herculano M.F. Alvarenga (MHNT), Celso Lira Ximenes (MUPHI), Hermínio Ismael de Araújo Júnior (UERJ), Cástor Cartelle (PUC Minas), Rafael Costa da Silva (MCT), Federico L. Agnolín (MACN), the late Storrs L. Olson (Smithsonian Institution), Darren Naish (University of Southampton), Ismar de Souza Carvalho (UFRJ), Graziella do Couto Ribeiro (MHNT), William Roberto Nava (MPM), Mariane Pereira Ferreira (MAE-USP/Scientia Consultoria Científica), Kamila L.N. Bandeira (MN), Levy Figuti (MAE-USP), Luiz Eduardo Anelli (IGc/USP), Thiago Vernaschi Vieira da Costa (Universidade Federal de Itajubá), Dione Seripierri (MZUSP), Igor Ferreira de Alvarenga, Helcio Alvarenga Júnior, Tereza Cristina Ferreira de Alvarenga, Marco Aurélio Crozariol (MHNC), Júlio Cesar de Almeida Marsola (Universidade Tecnológica Federal do Paraná), Melanie Nayyal (The Sixth Extinction Forum), Hélio Rosa de Miranda (MAE-USP), Gilberto Morais (MAE-USP), Juan M. Diederle (Universidad Autónoma de Entre Ríos), Kenneth E. Campbell, Jr. (LACM), Augusto S. Auler (Instituto do Carste), Alex Hubbe (Universidade Federal da Bahia), Mario Alberto Cozzuol (UFMG), Cécile Mourer-Chauviré (Université Lyon 1), Marcelo Adorna Fernandes (Universidade Federal de São Carlos), Hans Meltofte (Dansk Ornitologisk Forening), Laurent Raty (BirdForum), Björn Bergenholtz (BirdForum), Martin Schneider (BirdForum), Alexander Averianov (Russian Academy of Sciences), Matthew Shaw (Canterbury Museum), João Marcos (IPA), Luciana Barbosa de Carvalho (MN), Giovanna M.X. Paixão (Universidade Federal do Pampa), Alex Schiller Aires, Letícia Dutra Romualdo da Silva (MN), Geraldo Junior França, the staff of the Biblioteca Setorial Veríssimo de Melo of Museu Câmara Cascudo (UFRN), and two anonymous reviewers. Rafael also thanks José, Ivania, and Gabriela Nascimento, and Mariana Rodrigues, for their unconditional support. This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance Code 001. We also thank Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP, process 2017/23548-2) and Luís Fábio Silveira is supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico research productivity fellowship (process #302291/2015-6).
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How to cite this article
Nascimento RS, Silveira LF (2024) Fossil and subfossil birds of Brazil. Zoologia 41: e23079. https://doi.org/10.1590/S1984-4689.v41.e23079
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Published by
Sociedade Brasileira de Zoologia at Scientific Electronic Library Online (https://www.scielo.br/zool)
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[1]
Vidal and Campos ( 2008 ) mentioned birds among the material from the Romualdo Formation, possibly in error.
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[2]
Following the study’s Table 1 . There is some conflicting information regarding the feather’s classification in the study, with two semiplumes also regarded as contour feathers in some sections.
- [3]
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[4]
Campbell ( 1992: 20A) mentioned fossils of a giant Anhinga from the Amazon, without specifying the locality; even though he also mentioned fossils from Brazil (in Alvarenga 1992: 254), they may be the Anhinga fraileyi material he described in 1996. Campbell ( 1996: 2, 3) referred all the material described as A. fraileyi to Peru, with the holotype and referred material from the LACM 4611 locality, and the proximal end with the axis of a left humerus (LACM 135362) and the axis of a right humerus (LACM 135363) from the LACM 5158 locality. Diederle ( 2015: 132, 2017: 342) associated these two humeri fragments to the Brazilian territory, based on Negri et al. ( 2010: 258), who listed LACM 5158 as “Bandeira – Acre River (Brazil)”. Campbell ( 1996: 8) also mentioned that “a similar but less diverse paleofauna is known from Cachuela Bandeira, Bolivia, the second locality (LACM 5158) bringing bones of Anhinga fraileyi”. There is also the possibility that the mentions of Campbell (in Mourer-Chauviré [ 1989: 7]), Wall et al. ( 1991: 397), Alvarenga ( 1992: 254), Noriega ( 1992: 218), and Latrubesse et al. ( 1997: 112; they include Niterói and Patos as sources for the provided faunal list, without further details) refer to the A. fraileyi material described from Peru. Alvarenga and Höfling ( 2000: 589) mentioned Anhinga frailegyi [sic] as coming from the state of Acre, citing Campbell ( 1996 ) as a source. In that study’s second edition (Alvarenga and Höfling 2004: 827), they said the same for A. fraileyi. In the third edition (Alvarenga and Höfling 2011: 127), they mentioned A. fraileyi in Acre, citing both Campbell ( 1996 ) and Alvarenga and Guilherme ( 2003 ).
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[5]
Claussen published mentions of bird finds in 1841 (but read on), in the same work where he comments it was from the knowledge of William Buckland’s work in 1831 that he came up with the idea of looking for fossils in the Brazilian caves, finding mammalian remains the following year. Among the bird fossils he collected, we can highlight the material that served as a type for the description of Ciconia lydekkeri. However, by the date of his publication, it is not possible to determine the exact year in which these fossils were found, and Lund’s accounts are then the first documentations on the ancient remains of birds in Brazilian territory.
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[6]
According to Reinhardt ( 1881 ), this treatise, which should have been the concluding part of Lund’s fourth thesis (dated 30 January 1841 and sent for publication on the same occasion), was of moderate size, filling twenty pages in manuscript, with 13 colored illustrations of six bones supposedly belonging to five different bird species.
- [7]
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[8]
Opetiorrhynchus is replaced by Furnarius in Reinhardt ( 1881 ); Cypselus collaris, Capito melanotis, Coccyzus cajanus, and Crex minuta are synonyms, respectively, of Chætura zonaris (Shaw), Bucco chacuru (Vieillot), Pyrrhococcyx cayanus, and Porzana sp. e minimis in Winge’s work (Paula Couto 1950 ).
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[9]
“Family” erected by Illiger ( 1811 ) encompassing the dissimilar genera Glareola, Cereopsis, Dicholophus (=Cariama), Palamedea (=Anhima), Chauna, and Psophia. Reinhardt ( 1881 ) noted that Lund was not very aware of this group when he associated the remains of Rhea americana with the giant alectorid (see below).
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[10]
The rules established by Lund through the comparative study between the living and extinct mammals of the Lagoa Santa region include the concept that extinct fauna presented greater diversity and larger forms, which in the case of birds is exemplified by the alectorid. Truly giant extinct birds would only be discovered in South America in the late 1880s (Buffetaut 2013 ).
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[11]
Reinhardt ( 1881 ) noted that Lund’s and Gervais’ statements on Brazilian fossil birds were soon reproduced in several of the most important contemporary manuals and catalogs.
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[12]
Reinhardt reinforced this argument with a note in which he commented on the presence of unregistered bones of Chauna or Palamedea (=Anhima) in Lund’s collection, birds that do not occur regularly in the Lagoa Santa region, only occasionally, with an interval of several years. Winge ( 1887 ), however, noted that this material belongs to Chenalopex pugil, a species whose remains have until now been found only in this region.
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[13]
The official publication date for the first volume of “E Museo Lundii” is 1888. However, in a note accompanying the work’s index it appears: “O. Winge’s dissertation left the printer and was made available to the author as an offprint in October 1887; H. Winge’s in December of the same year, S. Hansen’s in September 1888”. This is the cause of much divergence observed on the date attributed to the works of Oluf and Herluf Winge. The review of Winge’s work published by Schäff (1888) in the Journal für Ornithologie is dated January 1888, and a small note about the work was published on the “Notices of Recent Ornithological Publications” of April 1888 by the editors of The Ibis (Sclater and Saunders 1888 ), which states its publication date is 1887. These publications support an 1887 date for the distribution of the offprint version of the work, and so it will be used here. Brodkorb ( 1964 ) cited it as “1887 (Oct. 1)”, probably a mistake from the original in Danish “…i Oktbr. 1887”.
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[14]
Lund divided the Brazilian vegetation based on his knowledge into “Matto Virgem” (virgin forest) and “Campos” (fields). The term “Campos”, in the sense used by Lund, is not used today in Brazil but was also recurrent in the works of Reinhardt and Eugenius Warming (Holten and Sterll 2011 ).
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[15]
Sick ( 1997 ) stated that Winge’s work was translated into Portuguese by Guttorm Hanssen. However, we were not able to find this translation.
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[16]
A German summary of Winge’s work was published by Schäff ( 1888 ).
- [17]
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[18]
Prado et al. ( 2016b ) mistakenly mentioned it being assigned to passerine birds in Martill and Filgueira’s study.
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[19]
Deposited in the MN collection according to Kellner ( 2006 ).
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[20]
Adapted from the plate’s caption. In the text, Naish et al. ( 2007: 526, 528) mentioned it as a “feather, associated with two or three smaller feathers and with some indeterminate bones” or “three asymmetrical feathers are preserved in association with several presumed carpal bones”.
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[21]
Candeiro ( 2015: 165) mistakenly described it as front limb phalanx.
- [22]
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[23]
This study includes an annex of the final manuscript of this analysis, authored by Taranto, Bergqvist, Loguercio, and Rocha-Barbosa, in the form submitted to the journal Acta Paleontologica Polonica. However, apparently it was not published.
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[24]
Bourdon et al. ( 2009 ) mistakenly attributed this record to Diogenornis fragilis.
- [25]
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[26]
Winge ( 1887: 11) used the term “Forskjellige Huler” (“various caves”) for what Lund seemed to refer to the Lapas da Anna Felicia, dos Coxos, dos Ossinhos, and da Serra das Abelhas. The surfaces and fractures of the bones are similar to those observed in the material from Lapa do Capão Seco.
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[27]
“bageste Haandrodsben” (“posterior carpal”) in the original.
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[28]
Winge ( 1887: 12) used the term “Fra nyeste Tid” (“of recent time”) associated with younger bones, often with dry tendons, found in the caves’ floors. Most were carried there by owls, but they may have also belonged to individuals who took shelter or were lost in these places. Skulls and other fragile parts are generally well preserved. Winge noted that the species represented in the collection only by these more recent bones were not included in his general list. Lambrecht’s ( 1933 ) “prähistorisch”.
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[29]
According to a review by J.J. (in McEvey [ 1968 ]).
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[30]
In the appendix where he lists the species that had remains of recent age found in the caves, Winge ( 1887: 50, 51) appears to have confused (or did not revise them after adding) the numbers of the species in his main list that he referenced in the description of the materials of “Phaëthornis pretrei?”. He mentioned the other trochilid discovered as No. 91 and not No. 92, and “Picid. g. sp., and minimis” he referenced “Picus (Campias) maculifrons” as No. 107 and not No. 108.
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[31]
This undergraduate thesis was not seen by us.
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[32]
Hsiou ( 2009a ) depicted the elements and attributed the numbers LGP-A0002 to a cervical vertebra and LGP-A0004 to the tibiotarsus.
- [33]
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[34]
Based on the fact that it is a humerus from the same locality, with preserved proximal end, and larger than that observed in Anhinga anhinga (the provided measurements are approximate). Several mentions of fossil anhingids from Acre have been made in the literature prior to Alvarenga and Guilherme ( 2003 ), but without specifications. These mentions are present, with question marks, in the synonyms of the records reported here.
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[35]
We were not able to find this site’s location. It is not the site with the same name in the municipality of Lassance, Minas Gerais, about 260 km from Lagoa Santa.
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[36]
Labeled as MN 3226-V in Patusco et al. ( 2016b ).
- [37]
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[38]
Pulsatrix koeniswaldiana Bertoni & Bertoni was considered close to Pulsatrix melanota (Tschudi), sometimes treated as conspecific. Because of this, Winge may have elected P. melanota as a possibility, even though the species, strictly speaking, is found only in Colombia, Ecuador, Peru, and Bolivia (Holt et al. 2017 ).
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[39]
In the main text in Alvarenga et al. ( 2011 ), the associated identification is DGM-1014, but, in the figures, it is that of the holotype skeleton, DGM-1418-R.
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[40]
Mesomyodes (or Clamatores), according to Mayr and Amadon ( 1951 ): Suboscines passeriforms in which the intrinsic muscles of the syringe, if present, are attached to one end or the middle of the bronchial semirings, as opposed to the Acromyodes, in which these muscles are attached at both ends.
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[41]
Grallaria (Grallaridae) previously included in Formicariidae (see Rice 2005 ).
Edited by
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Editorial responsibility:
Claudia Hermes
Publication Dates
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Publication in this collection
21 Oct 2024 -
Date of issue
2024
History
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Received
27 Oct 2023 -
Accepted
19 June 2024
