Rhea Brisson, 1760

Nascimento, Rafael S. & Silveira, Luís Fábio, 2024, Fossil and subfossil birds of Brazil, Zoologia (e 23079) 41, pp. 1-234 : 86-91

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. Quaternary – PI

Rhea – Guidon et al. 2009b: 80.

Rhea fossilis – Faure et al. 2010a: 23.

Rhea fossilis – Faure et al. 2010b: 5–7, fig. 8.

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).Howev- er, despite the rich vertebrate fauna excavated in the region since the 1980s, surprisingly no remains of these birds were found ( Guérin et al. 1993a, 1993 b, 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 celleci 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 Styltegaenger” – 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 Styltegaenger” (“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.

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 [25] Erroneously translated from “Løb” (“tarsometatarsus”) in Reinhardt (1881: 149) to “tibia” (“tibiotarsus”) in Reinhardt (1882: 329).

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. americanae [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, fig. 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, fig. 9. Rhea sp. [in part] – Jacobus and Schmitz 1983b: 269, fig. 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, fig. 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.

“avestruzes” – Goldmeier and Schmitz 1983: 22.

“emas” [?; in part?] – Prous 1992: 300.

“emas” [?; in part?] – Girelli and Rosa 2000: 60.

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.

Rhea americana – Mentz Ribeiro et al. 1989: 74.

Rhea americana – Mentz Ribeiro and Ribeiro 1999: 28.

Rhea americana – Rosa 2006f: 2.

cf. Rhea americana – Rosa 2009: 151.

Rheidae – Dias 2012: 15.

Rhea americana – Jacobus and Rosa 2013: 250.

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.

Rhea americana [in part] – Rosa 2001: C00011.

Rhea americana – Rosa 2006b: 231.

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

Late Holocene – RJ

Tinamidae [in part?] – Carvalho 1984: 56.

Tinamidae [in part] – Carvalho 1984: 122.

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.

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

Early Holocene – RS

Tinamidae [in part] – Rosa 2006f: 2.

Tinamidae indet. – Rosa 2009: 151.

Tinamidae [in part] – Dias 2012: 15.

Tinamidae [in part] – Hadler et al. 2013: 121.

Tinamidae [in part] – Jacobus and Rosa 2013: 250.

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.