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v m SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY ing is a list of referees who commented on one or more of the papers, and I am very grateful to all of them for their help: Antoni Alcover, S. Christopher Bennett, Zygmunt Bochenski, Alan Brush, Eric Buffetaut, Kenneth Campbell, Luis Chiappe, Charles Collins, Miguel Elorza, Andrzej Elzanowski, Steven Emslie, Per Ericson, Alan Feduccia, G.E. Goslow, Peter Houde, Helen James, Denes Janossy, Larry Martin, Gary Morgan, Cecile Mourer-Chauvire, Storrs Olson, John Ostrom, Kevin Padian, David Parris, Steven Parry, Stefan Peters, Gregory Pregill, J.H. Reichholf, Dale Serjeantson, David Steadman, Burkhard Stephan, Tommy Tyrberg, David Unwin, Kenneth Warheit, Paul Weldon, Peter Wellnhofer, Lawrence Witmer, Trevor Worthy, Zhonghe Zhou, and Richard Zusi. For assistance with transmission of manuscripts by electronic mail I am grateful to James Dean, Craig Ludwig, Chris Milensky, and Brian Schmidt. Mary Parrish repeatedly assisted with problems concerning illustrations and provided the outline key for the frontispiece. I also thank Sharon Jones, who cheerfully wielded her computer to render more readable several of the manuscripts that were more heavily scribbled by the editor. Storrs L. Olson

The Avifauna of Reunion Island (Mascarene Islands) at the Time of the Arrival of the First Europeans Cecile Mourer-Chauvire, Roger Bour, Sonia Ribes, and Francois Moutou ABSTRACT The excavations of five fossil bird localities on Reunion Island have yielded the remains of (1) five species still present on Reunion, which are mainly marine; (2) one extant species no longer on Reunion, the Greater Flamingo (Phoenicopterus ruber Linnaeus); and (3) 11 extinct species, namely, a night heron (Nycticorax duboisi (Rothschild)), an ibis {Threskiornis solitarius (Selys-Longchamps)), a sheldgoose (Alopochen kervazoi (Cowles)), a teal {Anas theodori Newton and Gadow), a falcon (Falco duboisi Cowles), a rail (Dryolimnas augusti, new species), a coot {Fulica newtonii Milne-Edwards), a pigeon (Nesoenas duboisi Rothschild), a parrot {Mascarinus mascarinus Linnaeus), an owl {Mascarenotus grucheti Mourer-Chauvire et al.), and a starling {Fregilupus varius (Boddaert)). Representatives of extinct endemic Mascarene taxa, such as the Raphidae, Aphanapteryx, Erythromachus, and large parrots of the genera Lophopsittacus and Necropsittacus, are so far unknown from Reunion. Except for Fulica newtonii, which probably colonized Reunion from Mauritius, and Dryolimnas augusti, all the other forms appear to have had normal or nearly normal flying ability. It is possible that Reunion was colonized by the ancestors of the same forms that colonized Mauritius and Rodrigues, but these forms may have been exterminated during the very explosive events of the last phase of volcanic activity of Piton des Neiges, which took place between 300,000 and 200,000 years ago. Reunion would then have been colonized again by forms that perhaps have not had enough time to lose their ability to fly. After the arrival of the first Europeans, all the larger endemic land birds became extinct, with the exception of Circus maillardi Verreaux. Although most of them were not morphologically flightless, and although the topography Cecile Mourer-Chauvire, Centre de Paleontologie Stratigraphique et Paleoecologie de I'Universite Claude Bernard-Lyon I, ERS 2042 du CNRS, 27-43 boulevard du II Novembre 1918, 69622 Villeurbanne Cedex, France. Roger Bour, Museum National d'Histoire Naturelle, Laboratoire des Reptiles et des Amphibiens, 25 rue Cuvier, 75005 Paris, France. Sonia Ribes, Museum d'Histoire Naturelle, 1 rue Poivre, 97400 Saint-Denis, La Reunion, France. Franqois Moutou, Minist'ere de I'Agriculture, CNEVA, Laboratoire Central de Recherches Veterinaires, 22 rue Pierre Curie, BP n° 67, 94703 Maisons-Alfort Cedex, France. of the island was very different, these birds became extinct on Reunion as rapidly as on the other Mascarene islands. Introduction The Mascarene Islands were "discovered" by Portuguese navigators as early as 1500 (North-Coombes, 1979). They were shown on more ancient maps of Arabian navigators but were uninhabited prior to the sixteenth century. Thus, Europeans were the first settlers to observe the very unusual fauna of these islands, one that was almost entirely exterminated within just two centuries. Numerous historical accounts of Reunion have been gathered into two publications by Albert Lougnon (1970, 1992), with the oldest account being that of Samuel Castleton, who landed on the island in 1613. The parts concerning the birds are given in extenso in the books of Barre and Barau (1982) and Barre et al. (1996). Particular points of these accounts have been discussed by Cheke (1987). The most complete and detailed report is that written by Dubois (1674), also known as Sieur D.B., who spent 16 months on the island in 1671 and 1672 (English translation by Oliver, 1897). These accounts are invaluable because they make it possible to follow the demise of the endemic fauna of Reunion from the beginning of the seventeenth century to the disappearance of the two most recently extinct species, Mascarinus mascarinus Linnaeus, in 1834, and Fregilupus varius (Boddaert), between 1838 and 1858. Fossil bird remains were found very early on Rodrigues (in 1786) and on Mauritius (in 1865), making it possible to have good knowledge of their vanished faunas. Until recently, however, the bird fauna of Reunion was known only from the accounts of early explorers. The first fossil remains were unearthed in 1974 by B. Kervazo during archaeological excavations in the Grotte des Premiers Francais. Subsequently, four other fossil sites containing bird remains also were discovered. The fossil birds of the present study came from these five

The Avifauna of Reunion Island (Mascarene Islands) at<br />

the Time of the Arrival of the First Europeans<br />

Cecile Mourer-Chauvire, Roger Bour, Sonia Ribes,<br />

and Francois Moutou<br />

ABSTRACT<br />

The excavations of five fossil bird localities on Reunion Island<br />

have yielded the remains of (1) five species still present on<br />

Reunion, which are mainly marine; (2) one extant species no<br />

longer on Reunion, the Greater Flamingo (Phoenicopterus ruber<br />

Linnaeus); and (3) 11 extinct species, namely, a night heron (Nycticorax<br />

duboisi (Rothschild)), an ibis {Threskiornis solitarius<br />

(Selys-<strong>Lo</strong>ngchamps)), a sheldgoose (Alopochen kervazoi<br />

(Cowles)), a teal {Anas theodori Newton and Gadow), a falcon<br />

(Falco duboisi Cowles), a rail (Dryolimnas augusti, new species),<br />

a coot {Fulica newtonii Milne-Edwards), a pigeon (Nesoenas<br />

duboisi Rothschild), a parrot {Mascarinus mascarinus Linnaeus),<br />

an owl {Mascarenotus grucheti Mourer-Chauvire et al.), and a<br />

starling {Fregilupus varius (Boddaert)). Representatives of extinct<br />

endemic Mascarene taxa, such as the Raphidae, Aphanapteryx,<br />

Erythromachus, and large parrots of the genera <strong>Lo</strong>phopsittacus<br />

and Necropsittacus, are so far unknown from Reunion. Except for<br />

Fulica newtonii, which probably colonized Reunion from Mauritius,<br />

and Dryolimnas augusti, all the other forms appear to have<br />

had normal or nearly normal flying ability. It is possible that<br />

Reunion was colonized by the ancestors of the same forms that<br />

colonized Mauritius and Rodrigues, but these forms may have<br />

been exterminated during the very explosive events of the last<br />

phase of volcanic activity of Piton des Neiges, which took place<br />

between 300,000 and 200,000 years ago. Reunion would then have<br />

been colonized again by forms that perhaps have not had enough<br />

time to lose their ability to fly. After the arrival of the first Europeans,<br />

all the larger endemic land birds became extinct, with the<br />

exception of Circus maillardi Verreaux. Although most of them<br />

were not morphologically flightless, and although the topography<br />

Cecile Mourer-Chauvire, Centre de Paleontologie Stratigraphique et<br />

Paleoecologie de I'Universite Claude Bernard-Lyon I, ERS 2042 du<br />

CNRS, 27-43 boulevard du II Novembre 1918, 69622 Villeurbanne<br />

Cedex, France. Roger Bour, Museum National d'Histoire Naturelle,<br />

Laboratoire des Reptiles et des Amphibiens, 25 rue Cuvier, 75005<br />

Paris, France. Sonia Ribes, Museum d'Histoire Naturelle, 1 rue<br />

Poivre, 97400 Saint-Denis, La Reunion, France. Franqois Moutou,<br />

Minist'ere de I'Agriculture, CNEVA, Laboratoire Central de Recherches<br />

Veterinaires, 22 rue Pierre Curie, BP n° 67, 94703 Maisons-Alfort<br />

Cedex, France.<br />

of the island was very different, these birds became extinct on<br />

Reunion as rapidly as on the other Mascarene islands.<br />

Introduction<br />

The Mascarene Islands were "discovered" by Portuguese<br />

navigators as early as 1500 (North-Coombes, 1979). They were<br />

shown on more ancient maps of Arabian navigators but were<br />

uninhabited prior to the sixteenth century. Thus, Europeans<br />

were the first settlers to observe the very unusual fauna of these<br />

islands, one that was almost entirely exterminated within just<br />

two centuries.<br />

Numerous historical accounts of Reunion have been gathered<br />

into two publications by Albert <strong>Lo</strong>ugnon (1970, 1992),<br />

with the oldest account being that of Samuel Castleton, who<br />

landed on the island in 1613. The parts concerning the birds are<br />

given in extenso in the books of Barre and Barau (1982) and<br />

Barre et al. (1996). Particular points of these accounts have<br />

been discussed by Cheke (1987). The most complete and detailed<br />

report is that written by Dubois (1674), also known as<br />

Sieur D.B., who spent 16 months on the island in 1671 and<br />

1672 (English translation by Oliver, 1897). These accounts are<br />

invaluable because they make it possible to follow the demise<br />

of the endemic fauna of Reunion from the beginning of the seventeenth<br />

century to the disappearance of the two most recently<br />

extinct species, Mascarinus mascarinus Linnaeus, in 1834, and<br />

Fregilupus varius (Boddaert), between 1838 and 1858.<br />

Fossil bird remains were found very early on Rodrigues (in<br />

1786) and on Mauritius (in 1865), making it possible to have<br />

good knowledge of their vanished faunas. Until recently, however,<br />

the bird fauna of Reunion was known only from the accounts<br />

of early explorers. The first fossil remains were unearthed<br />

in 1974 by B. Kervazo during archaeological<br />

excavations in the Grotte des Premiers Francais. Subsequently,<br />

four other fossil sites containing bird remains also were discovered.<br />

The fossil birds of the present study came from these five

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