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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unpublished measurements of Madagascan anatids.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-