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versión On-line ISSN 1851-8044

Ameghiniana vol.46 no.2 Buenos Aires abr./jun. 2009



Parrots (Aves, Psittaciformes) in the Pleistocene of Uruguay

Claudia P. Tambussi1, Carolina Acosta Hospitaleche1, Andrés Rinderknecht2 and Martín Ubilla3

1CONICET. División Paleontología Vertebrados, Museo de La Plata, Paseo del Bosque s/n, B1900FWA La Plata, Argentina.;
2Museo Nacional de Historia Natural y Antropología, Casilla de Correo 399, 11.000 Montevideo, Uruguay.
3Facultad de Ciencias, Universidad de la República, Iguá 4225, 11400 Montevideo, Uruguay.


Cyanoliseus Bonaparte, 1854 (Psittaciformes, Psittacidae, Arini) is the most frequent parrot in the Pleistocene record of Argentina. It comprises three species: the extinct C. ensenadensis (Cattoi, 1957) Tonni, 1972 and C. patagonopsis Acosta Hospitaleche and Tambussi, 2006, and the extant C. patagonus (Vieillot, 1817). A complete list of the Pleistocene Psittacidae known at the present is given by Acosta Hospitaleche and Tambussi (2006) and Tambussi et al. (2007). Nowadays, the burrowing parakeet Cyanoliseus patagonus (Psittacidae, Arini) is found in Argentina and the center of Chile, occasionally reaching Uruguay in winter (Collar, 1997; Bucher and Rodriguez, 1986) (figure 1). This paper reports the presence of Cyanoliseus in the Pleistocene of Uruguay, the first fossil parrot recorded in this country; comments about the environmental are also reported.

Figure 1. Location map showing the fossiliferous locality in Uruguay (A: Cyanoliseus ensenadensis; B: C. patagonus). Shaded area indicates the current area of ocurrence of Cyanoliseus patagonus in Uruguay / mapa de Uruguay señalando la ubicación de las localidades donde fueron exhumados los fósiles (A: Cyanoliseus ensenadensis; B: C. patagonus). El área sombreada indica el sector actual de distribución de Cyanoliseus patagonus.

Materials and methods

The material under study was compared with all living species inhabitant Uruguay and representatives of all genera of parrots living in Argentina (Appendix 1). Osteological terminology from Baumel and Witmer (1993) was used for the descriptions and their English equivalents through the discussion. Measurements are in millimeters with 0.1 mm accuracy.
The specimens studied are housed at the Museo Nacional de Historia Natural y Antropología, Montevideo, Uruguay (MNHN); Paleontología Vertebrados    of  the Facultad de Ciencias,
Universidad de la República, Uruguay (FC-DPV); Museo de La Plata, La Plata, Argentina (MLP); Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, Argentina (MACN).

Systematic paleontology

Order Psittaciformes Wagler, 1830
Family Psittacidae Illiger, 1811
Tribe Arini Smith, 1975

Genus Cyanoliseus Bonaparte, 1854

Type species. Cyanoliseus patagonus (Vieillot, 1817).

Remarks. The presence of the following associated characters is suitable to identify both humeri as Cyanoliseus: (1) deep and wide incisura capitis, (2) presence of a small tubercle in the center of the distal edge of the humerus head, (3) well developed crista deltopectoralis and (4) a deep incisura intercondylaris.
Comparative description. The distal extension of the crista deltopectoralis is larger than that of the crista bicipitalis, while in Brotogeris, Pionus, Pyrrhura, Forpus and Pionopsitta, both cristae are similar in their extension. As with other species of Cyanoliseus, the small tubercle in the center of the internal distal edge of the humerus head is present. The sulcus ligamentorum transversus is deeper and wider than in Amazona, Ara, Nandayus, Aratinga, Myiopsitta, Brotogeris, Pionus, Pyrrhura, Pionopsitta, Anodorhynchus and Forpus. The fossa musculus brachialis is deeper than in Ara, Aratinga, Amazona, Pionus, Pionopsitta, Pyrrhura, Myiopsitta, Forpus, Anodorhynchus and Brotogeris. The condylus dorsalis is elongated, while in Amazona, Ara, Aratinga, Myiopsitta, Brotogeris, Pionus, Pyrrhura, Pionopsitta, Anodorhynchus and Forpus, it is rounded. The condylus ventralis is oval and asymmetric, whereas in Amazona, Ara, Brotogeris, Pionus, Pyrrhura, Pionopsitta, and Forpus is symmetric.
The incisura intercondylaris is wider than in
Amazona, Ara, Nandayus, Aratinga, Myiopsitta, Brotogeris, Pionus, Pyrrhura, Pionopsitta, Anodor-hynchus and Forpus, and the incisura capitis is wider and deeper than in Amazona, Ara, Nandayus, Aratinga Brotogeris, Pionus, Pyrrhura, Pionopsitta, Anodorhynchus and Forpus.

Cyanoliseus patagonus (Vieillot, 1817) Figure 2

Figure 2. Humerus of Cyanoliseus patagonus MNHN 1716. 1, cranial view; 2, caudal view / húmero de Cyanoliseus patagonus MNHN 1716. 1, vista cranial; 2, vista caudal. cb, crista bicipitale; cd, condylus dorsalis; cdp, crista deltopectoralis; cv, condylus ventralis; ic, incisura capitis; sl, sulcus ligamentorus transversus; tb, tuberculum dorsale; tsv, tuberculum supracondylare ventrale.

Material. Complete right humerus (MNHN 1716).

Geographic locality. Department of Canelones, Balneario San Luis (Interbalnearia Route, km 64.5), Uruguay; coastal cliffs near the Río de la Plata estua-ry (34º 47' S; 55º 37' W) (figure 1).
Stratigraphic provenance. Libertad Formation (Pleistocene).
The fossil comes from a stratigraphic level composed by quartz-feldspar sands (reworked loess) at 90 cm under a horizon of calcretes (paleosoil). In this place the sediments -referred to the Libertad Formation- are composed of loess, reworked loess, brownish siltstone, paleosoils, and fluvial deposits (Loureiro et al., 2002). Further details about the age of these sediments are not possible at this stage. However, on the basis of its stratigraphic relation-ships, some authors have considered to be lower to middle Pleistocene in age (Bossi and Navarro, 1991; Panario and Gutiérrez, 1999).
Comments. According to the compared material, the
humerus length of living C. patagonus ranges from 47 to 51 mm (n=8). The length of the humerus of the MNHN-1716 specimen is included into the confidence limits of the mean of C. patagonus which indicates a non significant statistical difference between the fossil remains and the available sample for this character. Its size is intermediate between C. patagonopsis and C. ensenadensis.
Some differences allow us to differentiate it from C. patagonopsis: the tuberculum dorsale is proportionally bigger, the condylus dorsalis is more rounded and its edges are less acute, the condylus ventralis is more asymmetric, the tuberculum supracondylare ventrale is less robust, and the sulcus ligamentorum transversus is not so expanded.

Cyanoliseus ensenadensis (Cattoi, 1957) Figure 3

Figure 3. Humerus of Cyanoliseus ensenadensis FC-DPV-1417. 1, cranial view; 2, caudal view / húmero de Cyanoliseus ensenadensis FC-DPV-1417. 1, vista cranial; 2, vista caudal.

Material. Right humerus without distal portion (FC-DPV-1417).
Geographic locality. Sopas creek, Salto Department (31º15´S 57º00´W), Uruguay (figure 1).
Stratigraphic provenance. Brownish mudstones of the Sopas Formation (late Pleistocene). This unit is widespread in northern Uruguay and is in general characterized by massive brownish mudstones, sandstones and conglomerates formed mainly in a fluvial environment, but also including paleosoils (Ubilla et al., 2004).
Comments. C. ensenadensis is the smallest species of
the genus Cyanoliseus. The studied humerus is smaller than C. patagonus. The holotype, until now the only known specimen, measures 44.7 mm length. Although the sulcus ligamentorum transversus is small in this genus, it is even less expanded in C. patagonopsis and C. patagonus.

Discussion and conclusions

The Pampean Region of Cyanoliseus indicates that the three species lived during the Pleistocene, but only the two smallest species have been found in Uruguay: C. ensenadensis and C. patagonus.
Coincidently, all fossil species of Cyanoliseus are diagnosed using humeri, which are useful bones in the identification of these taxa. The humeral characters are: 1- a distal extension of the deltopectoral crest bigger than the bicipital crest, 2- a small tubercle in the center of the internal distal edge of the humerus head, 3- a deep and wide transverse ligamentary groove, 4- a deep brachial fossa, 5- an oval and asymmetric ventral condyle, 6- an enlarged dorsal condyle with its proximal end more acute, 7- a wide and deep capital groove, and 8- a broad dorsal tubercle.
The MNHN 1716 is morphologically indistinguishable from that of modern species. Cyanoliseus patagonus is a winter migrant whose presence in Uruguay was confirmed at the end of the 70's (Cuello, 1985; Azpiroz, 2001), and is recorded today in southwest of Uruguay, but only sporadically (Claramunt and Cuello, 2004) (figure 1).
The associated paleofauna with Cyanoliseus patagonus recovered from Libertad Formation at San Luis locality includes passerine birds, colubrids, and anurans (Rinderknecht, 1998, Claramunt and Rinderknecht, 2005), a woodpecker of the genus Colaptes Vigors, 1826 Rinderknecht and Claramunt, 2000), rodents of the genera Akodon Meyen, 1833 and Galea Meyen, 1831, and an assemblage of typical Pleistocene mammals (Ubilla and Rinderknecht, 2001). The cooccurrencee of the likely grazer Glyptodon Owen, 1839, the grass mouse Akodon, the cui Galea and this parrot supports, the idea of the presence of open or semiopen environments, considering the environmental adaptations inferred or known for these taxa (Rood, 1972; Walker, 1975; Scillato-Yané et al., 1995; Fariña and Vizcaíno, 2001). Living burrowing parrots almost always nest in holes (Masello and Quillfeldt, 2002), in the banks of rivers and other steep. An analysis related to environmental variables that could constrain the distribution of living representatives of C. patagonus, indicates subhumid to semiarid climates associated with 600-800 mm of annual precipitation (Tambussi et al., 2007).
The other fossil parrot described here belongs to
C. ensenadensis, characterized by its size and the transversal ligament groove which is less expanded than that of C. patagonopsis. Cyanoliseus ensenadensis is the smallest species of the genus, being a third smaller than the living species C. patagonus and two thirds smaller than the extinct C. patagonopsis (Acosta Hospitaleche and Tambussi, 2006).
Fauna associated with the humerus FC-DPV-1417 attributed to C. ensenadensis includes a variety of fossil vertebrates such as glyptodonts, horses like Equus neogeus and Hippidion principale, Machrauchenia patachonica, Toxodon platensis, deer like Ozotoceros, mostly related to open or semiopen environments (Ubilla, 2004; Ubilla et al., 2004) with a TL/OSL age of 43.5 +/-3.6 ka BP (Ubilla, 2004). Earthworm aestivation chambers described from this layer indicates a seasonal climate (Verde et al., 2007). In the Sopas Formation birds were found from different outcrops: aff. Rhea sp. Brisson, 1760, Chloephaga picta (Gmelin, 1789) and Cariama cristata (Linnaeus, 1766) (Ubilla et al., 2004; Tambussi et al., 2005). Open to semiopen environment under seasonal climatic conditions are suggested by paleoicnological evidence (Verde et al., 2007) and the record of seriemas and magellan gooses. According to mammalian biostratigraphic information, the faunal assemblage is correlated to the Lujanian Age/Stage of the Buenos Aires province, and 14C ages are >43 ky BP (Ubilla et al., 2004), which is older than the fauna of the Guerrero Member of the Lujan Formation (between 21ka and 10 ka BP, Cione and Tonni, 1999; Tonni et. al., 2003) of the Pampean Region.
In Argentina, Cyanoliseus ensenadensis has been recorded at Olivos (Ensenadan Formation) instead assignable to the Ensenadan stage (early to middle Pleistocene). Biostratigraphically, they are defined by
the presence of fossil mammals such as Mesotherium cristatum Serrés, 1867, Arctotherium angustidens Gervais and Ameghino, 1880, Theriodictis platensis Mercerat, 1891, Glyptodon munizi Ameghino, 1881, Neosclerocalyptus pseudornatus (Ameghino, 1889) Paula Couto, 1957 and Eutatus seguini Gervais, 1867. Taking into account biostratigraphic and absolute ages available (Ubilla, 2004), the record of C. ensenadensis in Uruguay belongs to later sediments than those of Argentina.
Until now, the fossil record of Cyanoliseus was limited to the Pleistocene of Argentina. Consequently, the material reported here not only expands the geographic range of Cyanoliseus during the Pleistocene, but is also the first time that any fossil Psittacidae has been reported from Uruguay. Therefore, the record of these parrots increases our knowledge of the Uruguayan Pleistocene avifauna, where Psittaciformes have never previously been found.

Appendix I. List of extant species and materials used for compar-isons. Fossil specimens are indicated with # / lista de especies y ma­teriales utilizados para las comparaciones. Los especimenes fósiles están indicados con #.

Amazona aestiva (Linnaeus, 1758) MLP 621, MACN 27741, 17308; Ara chloropterus Gray, 1859 MACN 53522, 20986; Ara militaris (Linnaeus, 1766) MLP 408; Aratinga acuticaudata (Vieillot, 1818) MLP 68; Aratinga aurea (Gmelin, 1788) MLP 342, MACN 54878; Aratinga leucophtalma (Statius Muller, 1776) MLP 407, MACN 20404; Anodorhynchus hyacinthinus (Latham, 1790) MACN 23591, 23590, Brotogeris versiculus MLP 348, MACN 18238; Cyanoliseus patagonus (Vieillot, 1817) MLP 65, MLP 260 a-g (n=8) and #MLP 81-VII-20-21, C. ensenadensis #MACN 17716, C. patagonopsis #MLP 81-VII-20-20, Forpus passerinus (Linnaeus, 1758) MACN 54856, Myiopsitta monachus Boddaert, 1783 MLP 66, 262, Nandayus nenday (Vieillot, 1823) MLP 257, 405, 406, Pionopsitta pileata (Scopoli, 1769) MLP 67, Pionus maximiliani (Kuhl, 1820) MLP 343, MACN 17309, Pyrrhura frontalis (Vieillot, 1818) MLP 351, MACN 17307.


To Santiago Claramunt who help us to study the materials from Uruguay. P. Tubaro (MACN) who facilitated comparisons with the extant parrots of the MACN under his care. D. Waterhouse and and anonymous reviewer provided helpful comments. It is a contribution to PICT 32617 (C. Tambussi), CSIC Proyect (M. Ubilla) and IGCP-518 (Bridgland-Latrubesse).


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Recibido: 24 de junio de 2008.
25 de marzo de 2009.

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