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localities, all of them situated in the territory of Saint-Paul, on the northwest portion of the island (Figure 1). The caves are not karstic cavities but are developed inside the lava flows, or at the junction between successive lava flows. ACKNOWLEDGMENTS.—We thank the Conseil general de la Reunion, the Conseil regional de la Reunion, and the Societe Reunionaise des Amis du Museum, whose generous cooperation made it possible to carry out excavations in the Marais de l'Ermitage, and we thank all the friends who helped us with these excavations and who provided material from other localities, in particular, Jean Jacques Argoud, Pierre Brial, Jean- Pierre and Colette Esmiol, Jean-Michel Probst, and the members of the Societe d'Etudes scientifiques des Cavernes de la Reunion. The work at the Smithsonian Institution was made possible by a Short Term Visitor Grant from the Office of Fellowships and Grants. For the loan of comparative modern and fossil material we thank Daniel Goujet and Christine Lefevre (Museum National d'Histoire Naturelle, Paris), Storrs L. Olson, (National Museum of Natural History, Smithsonian Institution), Robert Prys-Jones (The Natural History Museum, London), and Janet Hinshaw (University of Michigan). We thank Alain Dubois for his advice concerning nomenclatural problems and our two referees, Storrs Olson and David Steadman, for their very constructive criticism. Steve Goodman provided unpublished measurements of Madagascan anatids. For the xray pictures of the two specimens of Mascarinus mascarinus, we thank Jean Dorst, Paris, and Herbert Schifter, Vienna. The photographs are by Noel Podevigne, and the drawings are by Arlette Armand (Centre des Sciences de la Terre, Universite Claude Bernard-Lyon 1). ABBREVIATIONS AND ACRONYMS.—The following museum acronyms are used: AMNH, American Museum of Natural History, New York; BMNH, The Natural History Museum, London (formerly, British Museum (Natural History)); FSL, Faculte des Sciences de Lyon; LAC, Laboratoire d'Anatomie Comparee du Museum National d'Histoire Naturelle, Paris; MNHN, Museum National d'Histoire Naturelle, Paris; MHNR, Museum d'Histoire Naturelle, Saint-Denis, La Reunion; UCB, Universite Claude Bernard-Lyon 1; USNM, collections of the former United States National Museum, now in the National Museum of Natural History, Washington, D.C. In the listing of material, the following abbreviations are used: d., distal; j., juvenile; 1., left; p., proximal; r., right; s., shaft. Fossil Localities Grotte des Premiers Francais (Grande Caverne).—This is a very large cavity, situated 1.5 km from the center of Saint-Paul. Details concerning the stratigraphy and the location of excavations have been given by Kervazo (1979) and Bour (1979, 1980a). The bird material comes from layers 4 and 5 of pit number 2, in the most northeastern cave, which now includes a statue of the Virgin Mary. The top of layer 4 was situated about SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 80 cm below the floor of the cave at the time of the excavation, and layers 4 and 5 together were 25 cm thick. These layers did not include any trace of human occupation or any remains of introduced mammals; therefore, the vertebrate material can be considered as having been deposited prior to the occupation of the island by humans. In this cave, as well as in Grotte "au sable," most of the bird remains belong to the two species of shearwater, Puffinus pacificus Gmelin and P. Iherminieri Lesson. The remains come from all growth stages, from very young individuals, the bones of which are simple sticks without articulations, to fully grown adults. With the exception of a single bone of Fregilupus varius, no passerine remains were found. The sediments were not sieved (Kervazo, pers. comm., 1996), but a large number of very small bones, such as pedal phalanges of shearwaters, were collected, and, if there had been passerine remains, they would have been collected. The shearwater remains probably come from individuals that were nesting in burrows in the vicinity of the cave; the sediments of their nesting site might have been washed into the cave during a cyclonic episode. The other vertebrate remains probably come from animals that took refuge in the cave during such an episode (Bour, 1979). Bird material from this site was first described by Cowles (1987, 1994). All this material is in the MNHN and has the prefix LAC. Grotte de l'Autel.—This cave was discovered in 1980 by R. Bour and F. Moutou (Bour, 1980a). It is situated along the Nl road, 2 km south of Saint-Gilles, to the south of and a little higher up than a shrine dedicated to Saint Expedit. It is a small cavity, about 2.50 m long, 1.20 m wide, and 1 m high at its highest part. The sediment, completely removed in 1980, was not sieved. Grotte "au sable."—This cave also was discovered and excavated in 1980 by R. Bour and F. Moutou (Bour, 1980a); it was excavated again in 1987 by F. Moutou, R. Mourer, and C Mourer-Chauvire. It is situated close beside the Nl road, to the north and at the same level as the Saint Expedit shrine. Its dimensions are about 2 m 2 and 1.50 m high at its highest part. Its sediment also was completely excavated in 1987 and was sieved with 1.5 mm mesh screens. Very tiny lizard bones were found, which are smaller than those of recent endemic passerines, as well as isolated mouse teeth. It is therefore likely that the absence of passerines actually reflects their absence in the deposits and is not an artifact of collecting. The vertebrate remains from Grotte de l'Autel and Grotte "au sable" are probably contemporaneous with the occupation of the caves by the first settlers, from the middle of the seventeenth century, because they include some bones of introduced mammals. All the material from Grotte de l'Autel and Grotte "au sable" is in the UCB and has the prefix FSL. Marais de l'Ermitage.—This swamp is situated a little further to the south of the above-mentioned sites, along the Nl road, between Saint-Gilles and La Saline-les-Bains, at the locality of l'Ermitage-les-Bains. The first fossil bones were dis-
NUMBER 89 LePort St- Paul Grotte des premiers frangais St-G Mies' Grotte "au sable" Grotte de lAutel Marais de l'Ermitage Caverne de la Tortue 55°20 EST 55°40 EST St-Pierre FIGURE 1.—Map of Reunion Island showing the five localities in the northwest where fossil birds have been recovered. Contour intervals are in meters. covered in 1989-1990, during earthworks for the construction of the Jardin d'Eden. Excavations have been carried out in this same swamp, to the south of the Jardin d'Eden, under the responsibility of the Museum d'Histoire Naturelle de La Reunion, since 1992, and are still conducted every year. The area excavated so far is about 80 m 2 The filling of the swamp consists of about 80 cm of organic soil with very few fossils. This overlies a layer 30 to 40 cm deep made up of countless bones and shell fragments of the large, extinct land-tortoise Cylindraspis borbonica Bour, mixed with volcanic rocks and blocks of coral. Among this extraordinary accumulation of tortoise remains occur very rare bird and bat bones. The fossiliferous layer rests on marine sediments made up of coralline sands, fragments of corals, and marine molluscs. A very few remains of domestic mammals, introduced by humans, have been found associated with the extinct tortoises, al 21°SUD 21° 20 though most remains of domestic mammals are found in the upper sediments. The sediments of the marsh were systematically sieved using 2.5 mm and 1.5 mm mesh screens, and very small bones of juvenile Cylindraspis borbonica as well as tiny bones of bats were found. Here also, the absence of passerines probably reflects their absence in the sediments. Radiocarbon dates have been obtained for material from the Marais de l'Ermitage. The first one, on several bones of Cylindraspis borbonica, yielded an age of 915± 120 BP (Lyon 5551); interval in real years after calibration: 883-1273 AD (confidence interval 90%). Two other dates on single bones by the accelerator (AMS) method, gave the following results: Bone of Cylindraspis borbonica, from the base of the filling: 1755 ±40 BP (OxA-5994 (Lyon 201)). Interval in real years after calibration: 186-391 AD (confidence interval 95%).
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NUMBER 89<br />
LePort<br />
St- Paul<br />
Grotte des premiers frangais<br />
St-G Mies'<br />
Grotte "au sable"<br />
Grotte de lAutel<br />
Marais de l'Ermitage<br />
Caverne de la Tortue<br />
55°20 EST 55°40 EST<br />
St-Pierre<br />
FIGURE 1.—Map of Reunion Island showing the five localities in the northwest where fossil birds have been<br />
recovered. Contour intervals are in meters.<br />
covered in 1989-1990, during earthworks for the construction<br />
of the Jardin d'Eden. Excavations have been carried out in<br />
this same swamp, to the south of the Jardin d'Eden, under the<br />
responsibility of the Museum d'Histoire Naturelle de La<br />
Reunion, since 1992, and are still conducted every year. The<br />
area excavated so far is about 80 m 2 The filling of the swamp<br />
consists of about 80 cm of organic soil with very few fossils.<br />
This overlies a layer 30 to 40 cm deep made up of countless<br />
bones and shell fragments of the large, extinct land-tortoise<br />
Cylindraspis borbonica Bour, mixed with volcanic rocks and<br />
blocks of coral. Among this extraordinary accumulation of<br />
tortoise remains occur very rare bird and bat bones. The fossiliferous<br />
layer rests on marine sediments made up of coralline<br />
sands, fragments of corals, and marine molluscs. A very<br />
few remains of domestic mammals, introduced by humans,<br />
have been found associated with the extinct tortoises, al<br />
21°SUD<br />
21° 20<br />
though most remains of domestic mammals are found in the<br />
upper sediments.<br />
The sediments of the marsh were systematically sieved using<br />
2.5 mm and 1.5 mm mesh screens, and very small bones of juvenile<br />
Cylindraspis borbonica as well as tiny bones of bats<br />
were found. Here also, the absence of passerines probably reflects<br />
their absence in the sediments.<br />
Radiocarbon dates have been obtained for material from the<br />
Marais de l'Ermitage. The first one, on several bones of Cylindraspis<br />
borbonica, yielded an age of 915± 120 BP (Lyon<br />
5551); interval in real years after calibration: 883-1273 AD<br />
(confidence interval 90%). Two other dates on single bones by<br />
the accelerator (AMS) method, gave the following results:<br />
Bone of Cylindraspis borbonica, from the base of the filling:<br />
1755 ±40 BP (OxA-5994 (Lyon 201)). Interval in real years after<br />
calibration: 186-391 AD (confidence interval 95%).
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