Paläontologische Zeitschrift

, Volume 88, Issue 3, pp 353–359 | Cite as

The parrots (Aves: Psittaciformes) from the Middle Miocene of Sansan (Gers, Southern France)

ShortCommunication

Abstract

The occurrence of Archaeopsittacus sp. (Psittaciformes) in the fossil deposits of Sansan (France) is reported, testifying to a survival of this genus in the Middle Miocene, as Archaeopsittacus verreauxi was described from the Early Miocene of Saint-Gérand-le-Puy (France) and was recorded only from its type locality. The data discussed here indicate the presence of two parrot species from Sansan, as it is the type locality of another parrot species, Pararallus dispar, only known from this locality. The differences between the humeri of these two taxa are described in detail, together with the differences from the other European fossil parrot species. The presence of more than one species of parrot in the same locality is not rare, but in Europe it is recorded in Sansan for the second time. Evidence for parrots also confirms the paleoenvironmental reconstruction of Sansan, as parrots are primarily arboreal species. Archaeopsittacus also represents one of the few common elements between the Early and Middle Miocene European bird assemblages.

Keywords

Aves Psittaciformes Miocene France 

Kurzfassung

Das Vorkommen von Archaeopsittacus sp. (Psittaciformes) im Fossilbericht von Sansan (Frankreich) belegt das Überleben dieser Gattung bis zum Mittleren Miozän, da Archaeopsittacus verreauxi zuvor nur von der Typuslokalität Saint-Gérand-le-Puy (Frankreich) aus dem frühen Miozän bekannt war. Sansan ist auch die Typuslokalität und einziger Fundort für eine weitere Papageienart, Pararallus dispar. Merkmale des Oberarmknochens, die eine Identifizierung beider Arten und ihre Unterscheidung von anderen Papageien aus dem Fossilbericht Europas zulassen, werden detailliert beschrieben. Das Vorkommen von mehr als einer Papageienart an derselben Fossilfundselle ist nicht selten, aber Sansan ist erst die zweite Fundstelle in Europa, an der sich mehr als eine Papageienart feststellen lässt. Da Papageien in ihrer Lebensweise in erster Linie arboreal sind, stützt der Fossilbeleg frühere Rekonstruktionen der Paläohabitate von Sansan. Archaeopsittacus ist darüber hinaus eine der wenigen Taxa, die sowohl in früh- wie in mittelmiozänen Avifaunen vertreten sind.

Stichwörter

Aves Psittaciformes Miozän Frankreich 

Introduction

Fossil parrots are not a common finding, with some records from the Plio-Pleistocene of Africa (Manegold 2013 and reference therein), the Neogene and Pleistocene of America (Wetmore 1926; Campbell 1979; Tonni and Noriega 1996), and the Neogene of Australia and New Zealand (Boles 1993, 1998; Worthy et al. 2007, 2011). In Europe the parrots are reported from the Late Oligocene to Middle Miocene with five species described from various localities in France, Germany and Czech Republic (Mayr and Göhlich 2004; Mayr 2010). The various European species are, up to now, only known from their type localities, with all the species well characterized by their tarsometatarsi, known in all five taxa (Mayr and Göhlich 2004: fig. 1; Mayr 2010).

Specifically, Mogontiacopsitta miocaena was described on the basis of an incomplete tarsometatarsus from the Late Oligocene/Early Miocene of the Mainz Basin, Germany (Mayr 2010). In addition, from the same locality, two distal tibiotarsi of two different species of Psittacidae were reported, but none of them were certainly referred to Mogontiacopsitta miocaena (Mayr 2010). Archaeopsittacus verreauxi was described from the Early Miocene (MN 2) of Saint-Gerand-le-Puy, France, on the basis of a tarsometatarsus, a distal tibiotarsus and three humeri (Milne-Edwards 1867–1871). Xenopsitta fejfari was described from a tarsometatarsus and two fragmented humeri from the Early Miocene (MN 3) of Merkur, Czech Republic (Mlíkovský 1998). Bavaripsitta ballmanni was found in the Middle Miocene (MN 6) of Steinberg in the Nördlinger Ries, Germany, and is known by its tarsometatarsus and a tentatively referred distal humerus (Mayr and Göhlich 2004). Pararallus dispar is a small parrot from the Middle Miocene (MN 6) of Sansan, France, known by different isolated bones, including an incomplete tarsometatarsus (Cheneval 2000). This species was originally described as a species of Rallidae by Milne-Edwards (1867–1871), and the syntypical series includes the distal humerus, which was selected as the lectotype by Cracraft (1973: 33). According to Cheneval (2000) this humerus belongs to Psittacidae, while the other syntypical bones were referred to Palaeoaramides beaumontii (Rallidae). Milne-Edwards (1872: 1033) recognized, among the fossil bird bones from Sansan, the presence of a parrot smaller than Archaeopsittacus verreauxi, for which he proposed the name Psittacus Latertianus. However, according to Cheneval (2000) and Mayr and Göhlich (2004) and contra Mlíkovský (2002), this is a nomen nudum and cannot be applied to the material of Sansan instead of Pararallus dispar, as its selected lectotype cannot be changed without the permission of the International Commission of Zoological Nomenclature.

Following Mayr and Göhlich (2004), it is not possible to assign the various European taxa of Psittaciformes to any modern bird family or genus, or ever to an extinct one, because of the scarcity of available fossil material and for the incomplete understanding of the relationships between the various extant taxa.

Study of the fossil remains found in the Middle Miocene locality of Sansan revealed that among the bones previously referred to Pararallus dispar by Cheneval (2000), one humerus is attributable to Archaeopsittacus sp. by size and morphology. This finding testifies to the survival of the latter genus until the Middle Miocene and represents the second European record of two species of parrot in the same locality. In addition, morphological differences in the humeri of the two taxa are described in details.

Materials and methods

The fossil bones here studied are preserved in the collections of the Museum National d’Histoire Naturelle (MNHN), Paris, France. They were compared with recent bird skeletons stored in the MNHN and in the Institut de Palèontologie Humaine (IPH), Paris, France, in the Museo Civico di Storia Naturale di Carmagnola, Torino, Italy (MCCI), and in the Dipartimento di Scienze della Terra of the Torino University, Italy (Marco Pavia Ornithological Collection, MPOC). The osteological terminology follows Baumel and Witmer (1993).

Systematic paleontology

Order Psittaciformes Wagler 1830

Genus Pararallus Lambrecht 1933

Pararallus dispar (Milne-Edwards 1867–1871)

Figs. 1a–f; 2c, f
Fig. 1

Pararallus dispar from the Middle Miocene of Sansan, Gers, France (MN 6). a Left humerus, holotype (MNHN Sa 1201), cranial view; b right humerus (MNHN Sa 14104), caudal view; c left ulna (MNHN Sa 14102), cranial view; d right coracoid (MNHN Sa 14106), dorsal view; e left tibiotarsus (MNHN Sa 1606), cranial view; f left tarsometatarsus (MNHN Sa 1668), dorsal view. The scale bars represent 5 mm

Fig. 2

Humeri of fossil parrots in comparison. a, dArchaeopsittacus verreauxi, left humerus (paralectotype MNHN Av 2839), caudal (a) and cranial (d) views. b, eArchaeopsittacus sp., left humerus (MNHN Sa 1216), caudal (b) and cranial (e) views. c, fPararallus dispar, reversed right humerus (MNHN Sa 14104), caudal (c) and cranial (f) views. cvf crus ventral fossae, ch caput humeri, fo fossa olecrani, ih intumescentia humeri, mc margo caudalis, td tuberculum dorsale. The scale bars represent 5 mm

Synonyms

Rallus dispar Milne-Edwards 1867–1871:157 (vol. 2), pl. 105, figs. 23–26 (original description).

Rallus dispar Milne-Edwards 1867–1871:155–156 (vol. 2), pl. 105, figs. 17–22, 27–30 (non).

Psittacus lartetianus Milne-Edwards 1872: 6 (nomen nudum).

Rallus dispar Milne-Edwards 1867–1871: Lydekker 1891: 144 (in part).

Pararallus dispar (Milne-Edwards 1867–1871): Lambrecht 1933: 466 (new combination).

Pararallus dispar (Milne-Edwards 1867–1871): Brodkorb 1967: 120.

Pararallus dispar (Milne-Edwards 1867–1871): Cracraft 1973:33, fig. 15a–b (in part).

Pararallus dispar (Milne-Edwards 1867–1871): Mlíkovský 1998: 338.

Psittacus lartetianus (Milne-Edwards 1872): Mlíkovský 2002: 147.

Palaeoaramides beaumontii (Milne-Edwards 1867–1871): Mlíkovský 2002: 175 (in part).

Pararallus dispar (Milne-Edwards 1867–1871): Mayr and Göhlich 2004: 49, fig. 1E–F.

Pararallus dispar (Milne-Edwards 1867–1871): Cheneval 2000: 363, fig. 20.

Holotype Left humerus, distal half (MNHN Sa 1201) from Sansan, selected by Cracraft (1973: 33, fig. 15a–b).

Other topotypical material Two left coracoids, cranial fragments (MNHN Sa 14107, MNHN SA 14108); right coracoid (MNHN Sa 14106); left humerus (MNHN Sa 1274); two distal left humeri (MNHN Sa 1490, MNHN Sa 10282); two right humeri (MNHN Sa 1492, MNHN Sa 14104); two left ulnae (MNHN Sa 1464, MNHN Sa 14102); left ulna, proximal end (MNHN Sa 14103); left ulna, distal half (MNHN Sa 1314); left tibiotarsus, distal half (MNHN Sa 1606); left tarsometatarsus (MNHN Sa 1669); right tarsometatarsus (MNHN Sa 1668). Among the topotypical material listed by Cheneval (2000: 363) the left humerus MNHN Sa 1216 is belonging to Archaeopsittacus sp. (see below). A right ulna (MNHN Sa 1312) and a right radius (MNHN Sa 10230) are here considered not belonging to a Psittaciformes and have to be deleted from the series of Pararallus dispar.

Type locality and horizon Sansan, Gers, France; Middle Miocene, Mammal Neogene zone MN 6.

Measurements see Tables 1 and 2.
Table 1

Measurements (in mm) of the bones of Pararallus dispar from the Middle Miocene of Sansan, Gers, France (MN 6)

 

GL

Wp

Dp

Wd

Dd

Ws

Ds

Lm

Coracoid

  MNHN Sa 14106

4.4

2.7

2.1

2.2

1.7

19.7

  MNHN Sa 14107

4.0

2.6

2.1

1.8

  MNHN Sa 14108

4.1

2.6

 

GL

Wp

Dp

Wd

Dd

Ws

Ds

 

Humerus

  MNHN Sa 1201 (holotype)

5.7

3.9

2.2

2.5

 

  MNHN Sa 1274

23.7

2.8

5.4

3.7

(2.6)

 

  MNHN Sa 1490

5.5

3.6

3.0

2.5

 

  MNHN Sa 1492

26.5

8.5

4.5

5.7

3.9

3.1

2.5

 

  MNHN Sa 10282

5.7

3.9

2.8

2.5

 

  MNHN Sa 14104

26.8

9.6

4.9

5.8

4.1

3.0

2.6

 

Radius

  MNHN Sa 10230

(3.8)

2.5

1.8

1.7

 

Ulna

  MNHN Sa 1314

4.1

3.6

2.1

2.1

 

  MNHN Sa 1464

(32.0)

4.7

(3.7)

4.5

3.5

2.3

2.3

 

  MNHN Sa 14102

29.5

4.4

4.1

3.8

3.4

2.2

2.2

 

  MNHN Sa 14103

4.6

4.1

2.5

2.4

 

Tibiotarsus

  MNHN Sa 1606

3.9

3.2

1.6

1.6

 
 

GL

Wp

Dp

Wd

Dd

Ws

Ds

Dpm

Tarsometatarsus

  MNHN Sa 1668

3.7

2.0

1.3

2.3

  MNHN Sa 1669

4.2

2.1

1.3

2.4

Measurements of bones that are slightly worn or damaged are given in parentheses

GL, greatest length; Wp, proximal width; Dp, proximal depth; Wd, distal width; Dd, distal depth; Ws, smallest width of shaft. Wd, smallest depth of the shaft. Coracoid: Lm, medial length. Tarsometatarsus: Dpm, smallest proximal depth without the hypotarsus

Table 2

Measurements (in mm) of the complete humeri of Archaeopsittacus verreauxi, Archaeopsittacus sp., and Pararallus dispar from the Early Miocene of Saint-Gérand-le-Puy, Allier, France (MN 2), and from the Middle Miocene of Sansan, Gers, France (MN 6)

 

GL

Wp

Dp

Wd

Dd

Ws

Wd

Archaeopsittacus verreauxi

 MNHN Av 2839

38.7

11.7

6.8

8.3

5.2

4.2

3.8

 MNHN Av 2840

37.8

11.3

6.3

7.5

3.7

 MNHN Av 2841

37.0

11.5

5.8

7.8

5.4

4.1

3.7

Archaeopsittacus sp.

 MNHN Sa 1216

(33.5)

10.0

5.4

6.9

4.5

3.4

2.9

Pararallus dispar

 MNHN Sa 1274

23.7

2.8

5.4

3.7

(2.6)

 MNHN Sa 1492

26.5

8.5

4.5

5.7

3.9

3.1

2.5

 MNHN Sa 14104

26.8

9.6

4.9

5.8

4.1

3.0

2.6

Measurements of bones that are slightly worn or damaged are given in parentheses

GL, greatest length; Wp, proximal width; Dp, proximal depth; Wd, distal width; Dd, distal depth; Ws, smallest width of shaft. Wd, smallest depth of the shaft

Stratigraphical and geographical distributionPararallus dispar is only reported from the Middle Miocene (MN 6) of Sansan (Gers, France).

Remarks The various bones of this species were described by Cheneval (2000), even if he did not compare them directly with the other European fossil parrots. In more detail, the bones of Pararallus dispar are stout with the following morphological characteristics: the humerus shows a well-expanded crus dorsale fossae, the tuberculum dorsale is elongated and not separated from the caput humeri, the intumescentia humeri are squared, and a well-evident small tuberculum is on the distal part of the shaft, proximally to the condylus dorsalis. The coracoid shows the facies articularis clavicularis caudally extended and the cotyla scapularis with a caudal ridge in the coracoid; the ulna shows a well-marked tuberculum ligament collateralis ventralis and impressio brachialis; the tibiotarsus shows two tuberculi on the lateral and ventral side of the distal part of the shaft and the presence of the pons supratendineus on the distal end; the tarsometatarsus shows a protruding tuberositas musculi tibialis cranialis hypotarsus. Mayr and Göhlich (2004) and Mayr (2010) made detailed comparisons of the tarsometatarsi of the five European Psittaciformes to assess the differences among the various species on the basis of the single bone preserved in all of them. The detailed morphological analysis of the preserved bones of Pararallus dispar could give new information about its affinities among the various families within the Psittaciformes, but this is beyond the scope of this article.

Genus Archaeopsittacus Lambrecht 1933.

Archaeopsittacus sp.

Fig. 2b, e.

Referred Material Left humerus with a damaged proximal end (MNHN Sa 1216) from the Middle Miocene (MN 6) of Sansan (Gers, France).

Measurements see Table 2.

Remarks The humerus of Archaeopsittacus sp. is relatively stout and shows the typical features of Psittaciformes in respect of Columbiformes, with the fossa pneumotricipitalis relatively small, a well-marked impressio coracobranchialis, a step between the incisura capitis and the diaphysis, the condylus ventralis not protruding distally, the processus supracondylaris dorsalis absent, and the processus flexorius rounded and protruding caudally.

The humerus MNHN Sa 1216 is smaller than the corresponding bones of Archaeopsittacus verreauxi, but larger than those of Pararallus dispar (see Table 2); it shares some morphological characteristics with the paratypes of A. verreauxi. In particular, in respect to Pararallus dispar, the humerus MNHN Sa 1216 shows the crus ventral fossae less expanded ventrally, the tuberculum dorsale more rounded and well separated from the caput humeri, the margo caudalis proximally jointed with the caput humeri, the intumescentia humeri ventrally more rounded and the crista deltopectoralis more protruding cranially, the fossa olecrani well marked, and without the small tuberculum on the shaft, proximally to the condylus dorsale, well evident in Pararallus dispar.

The humeri of the other fossil European parrots can be excluded from comparison as they show different proportion in respect to MNHN Sa 1216. In particular, the humerus of Xenopsitta fejfari is stouter, according to the measurements given by Mlíkovský (1998: 336), and the humerus tentatively referred to Bavaripsitta ballmanni is clearly smaller (Mayr and Göhlich 2004). The humerus of Mogontiacopsitta miocaena is not known, but this species can be excluded as it shows a smaller and stouter tarsometatarsus than that of Archaeopsittacusverreauxi, also with some morphological differences (Mayr 2010).

The great similarity between the humerus from Sansan and the paralectotypical humeri of Archaeopsittacus verreauxi from Saint-Gérand-le-Puy, and the different proportion of the humeri of the other fossil species, allow referring the left humerus MNHN Sa 1216 to Archaeopsittacus sp. Hopefully new fossil material from Sansan or another coeval locality could clarify the relationships of this taxon of the Middle Miocene with A. verreauxi.

Discussion

The data presented here indicate the occurrence of two parrot species in the Middle Miocene of Sansan, where Cheneval (2000) only reported the presence of Pararallus dispar. Although the coexistence of more than one species of parrots in the same locality has been reported (Worthy et al. 2011; Manegold 2013), Sansan is only the second case for Europe. The fossil avifauna of Sansan was first studied by Milne-Edwards (1867–1871) and later revised and updated by Cheneval (2000). It is well diversified and includes taxa referable to ten modern bird orders; the avifauna is characterized by aquatic birds, in terms of the number of taxa and bones, but it also includes some land birds. The presence of two species of Psittaciformes reinforces the indication of arboreal formations already suggested by Cheneval (2000) in his paleoenvironmental reconstruction, as most parrots are primarily arboreal species using trees as a food resource or nesting location or both (del Hoyo et al. 1997).

The avifauna of Sansan is very different from that of Saint-Gérand-le-Puy in terms of species composition. In fact, Archaeopsittacus sp. is one of the few common elements within the two fossil bird associations (Cheneval 2000; Göhlich and Mourer-Chauviré 2005; Mourer-Chauviré et al. in press), testifying to the great change in the avifauna occurring in Europe from the Early Miocene to the Middle Miocene, especially in the aquatic taxa. However, the revision of the poorly represented taxa found in the various localities could probably reveal major similarities within European Miocene bird associations, as some species described from Sansan have already been reported from other European localities of the same age or even younger (Cheneval 2000; Pavia 2013).

Notes

Acknowledgments

R. Allain, C. Lefevre (MNHN), G. Boano (MCCI), and H. de Lumley (IPH) are thanked for the possibility to access the collections at their institutions, and A. Manegold and G. Mayr (Forschungsinstitut Senckenberg) for reviewing the manuscript and for the help with the German abstract. The work was financially supported by Italian MIUR PRIN 2009MSSS9L_002 to Giulio Pavia and by Synthesys projects FR–TAF–2607 and NL–TAF–225.

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Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  1. 1.Dipartimento di Scienze della Terra, Museo di Geologia e PaleontologiaUniversità degli Studi di TorinoTurinItaly

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