Colaptes Vigors, 1825

Nascimento, Rafael S. & Silveira, Luís Fábio, 2024, Fossil and subfossil birds of Brazil, Zoologia (e 23079) 41, pp. 1-234 : 162-170

Colaptes Vigors

544. Colaptes sp.

Early Holocene – GO

Colaptes sp. – Rosa 2004: 237.

Colaptes indet [?] – Paulo 2009: 142.

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.

Piciformes [in part] – Guérin et al. 1993a: 198.

Piciformes [in part] – Guérin et al. 1993b: 328.

Colaptes melanochloros – Guérin et al. 1996: 85.

Colaptes melanochloros – 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.

546. Colaptes cf. campestris (Vieillot)

Late Holocene – RS

Colaptes campestris – Rosa 2001: C00011.

Colaptes cf. campestris – Rosa 2006b: 232.

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. 31D

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, fig. 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, fig. 61. Paleopsilopterus itaboraiensis – Agnolín 2009b: 21, fig. 18. Paleopsilopterus itaboraiensis [in part] – Alvarenga et al. 2011: 191–192, fig. 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) comment- ed 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: 51 R. 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, fig. 7.2.

Alvarenga et al. (2011) tentatively attributed to this species five ungual phalanges collected by Rubens da Silva Santos during the 1970 s 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).

Paleopsilopterus itaboraiensis – Metello et al. 2012b: 140. Paleopsilopterus itaboraiensis [in part?] – Metello et al. 2014: 84. Paleopsilopterus itaboraiensis [in part] – Metello and Bergqvist 2014: 151.

Metello et al. (2012b) and Metello and Bergqvist (2014) associated with this species another ungual phalanx (MCT 1836-R), attributed to digit IV. Metello et al. (2012b) 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 assump - tion 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. 31C

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, fig. 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)

Late Holocene – MS

“aves médias (possivelmente seriemas)” [in part?] – Pacheco et al. 2007: 302.

“aves médias (possivelmente seriemas)” [in part?] – Pacheco and Martins 2009a: 172.

“aves média (seriemas)” [sic] – Pacheco 2008: 128.

“aves médias (seriemas)” – Pacheco and Martins 2009b: 154.

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

Middle Eocene – PR

Phorusrhacidae – Sedor et al. 2014a: 614.

Aves – Cunha et al. 2014: 691.

Phorusrhacidae – Sedor et al. 2014b: 807.

Phorusrhacidae – Cunha 2016: 68.

Phorusrhacidae – Sedor et al. 2016: 40.

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, fig. 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, fig. 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, fig. 31.9.

Paraphysornis brasiliensis – Alvarenga and Höfling 2003: 70, figs 1–4, 6, 15–17, 34.

Paraphysornis brasiliensis – Alvarenga and Höfling 2004: 828, fig. 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, fig. 7.11.

Paraphysornis brasiliensis – Angst and Buffetaut 2017: 137, fig. 5.9.F.

Paraphysornis brasiliensis – Angst and Chinsamy 2017: 17–37, fig. 1B.

Paraphysornis brasiliensis – Carmo et al. 2024: 5, fig. 3A–B.

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.?)

Late Holocene – RS

Falconiformes [?; in part] – Rosa 2001: C00011.

Falconiformes – Rosa 2006b: 231.

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.?)

Early Holocene – GO

Falconiformes indet. [in part] – Rosa 2004: 230.

Falconiformes sp. indet. [in part] – Rosa 2004: 233. Falconiformes sp. indet. [in part] – Rosa 2004: 239. Falconiformes indet. [in part] – Rosa 2004: 247.

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

Late Holocene – SC

Falconiformes – Schmitz and Ferrasso 2011: 147, fig. 5E. Falconiformes – Ferrasso and Schmitz 2013: 130.

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

Holocene – RN

“falconiformes” – Silva 2013: 91.

Falconiformes – Silva 2014: 133.

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

Early Holocene – RS

cf. Falconidae – Rosa 2006f: 2.

cf. Falconidae – Rosa 2009: 151.

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