Anhinga Brisson, 1760

Nascimento, Rafael S. & Silveira, Luís Fábio, 2024, Fossil and subfossil birds of Brazil, Zoologia (e 23079) 41, pp. 1-234 : 138-143

Anhinga Brisson

454. † Anhinga sp.

Late Miocene – AC

Fig. 25A–C

Anhinga indet. – Bandeira et al. 2015: 115.

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)

Late Holocene – RJ

Anhinga anhinga – 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 (“biguá, biguatinga”) [in part] – Magalhães et al. 2001: 59.

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

Late Miocene – AC

Fig. 25D–E

Anhinga cf. A. minuta – Bandeira et al. 2015: 115.

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 .

457. † Anhinga minuta Alvarenga & Guilherme

Late Miocene – AC

Fig. 34B

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, fig. 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, fig. 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

Late Miocene – AC

Anhingidae [in part] – Hsiou et al. 2022b: 15.

Macranhinga sp. – Guilherme et al. 2023: 5–7, fig. 2A–E.

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).

Anhingidae [in part] – Hsiou et al. 2022b: 15.

Macranhinga sp. indet. 2 (robust) – Guilherme et al. 2023: 8–9, fig. 3E, 3G, 3I, 3J.

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

Late Miocene – AC

Anhingidae [in part] – Hsiou et al. 2022b: 15.

Macranhinga sp. – Guilherme et al. 2023: 7–8, fig. 2F–H.

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

Late Miocene – AC

Anhingidae [in part] – Hsiou et al. 2022b: 15.

Macranhinga sp. indet. 1 (gracile) – Guilherme et al. 2023: 8, fig. 3A, 3C, 3J.

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

Late Miocene – AC

Anhingidae [in part] – Hsiou et al. 2022b: 15.

Macranhinga sp. – Guilherme et al. 2023: 9–10, fig. 4A, 4C.

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

Late Miocene – AC

Anhingidae [in part] – Hsiou et al. 2022b: 15.

Macranhinga sp. – Guilherme et al. 2023: 10, fig. 4E–F.

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. 34C

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, fig. 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, fig. 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.

Aves – Loboda et al. 2019: 136.

Macranhinga ranzii – Guilherme et al. 2020: 3091–3100, figs 2–5.

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.