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32 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Without any justification, Rothschild (1907) placed this form in the genus Necropsittacus of Rodrigues and described it as N. borbonicus. Family STRIGIDAE Genus Mascarenotus Mourer-Chauvire, Bour, Moutou, and Ribes Mascarenotus grucheti Mourer-Chauvire, Bour, Moutou, and Ribes, 1994 Gmchet's Mascarene Owl FIGURE 13^ MATERIAL.—Grotte des Premiers Francais. Holotype: r. tarsometatarsus, 1993-49. Paratypes: 1. humems, 1993-50; r. p. tibiotarsus, 1993-48; r. d. tibiotarsus, 1993-47. Grotte de l'Autel: r. tarsometatarsus, 330537; r. d. tarsometatarsus, 330538. Grotte "au sable": 1. quadrate, 330809; 1. p. femur, 330737. Marais de l'Ermitage: r. tarsometatarsus, 1800. REMARKS.—We have not found any additional remains other than those used in the original description of Mascarenotus grucheti (Mourer-Chauvire et al., 1994). We have placed all the Mascarene strigiforms in the extinct genus Mascarenotus, which resembles the genus Otus but presents some distinctive morphological characteristics. This genus includes one species on Mauritius, Mascarenotus sauzieri, based on Strix sauzieri Newton and Gadow, 1893 (synonyms, Otus commersoni Oustalet (1896), Strix newtoni Rothschild (1907)), one species on Rodrigues, Mascarenotus murivorus based on Strix {Athene) murivora Milne-Edwards (1874) (synonym, Bubo (?) leguati Rothschild (1907)), and one species on Reunion, Mascarenotus grucheti Mourer-Chauvire et al., 1994. The tarsometatarsi of M. grucheti are very close in size to those of M. sauzieri, although the only humems known from Reunion is clearly smaller than those of M. sauzieri, for which we have a good sample of comparative material (12 specimens). The ratio of element lengths (Mourer-Chauvire et al., 1994, fig. 1) shows M. sauzieri to be strikingly parallel to modem insular species of the genus Otus, such as Otus lawrencii (Sclater and Salvin), or O nudipes (Daudin), from the West Indies, in which the legs are much more elongated than in continental forms. The same adaptation is found in the four extinct insular species of the genus Grallistrix, from Hawaii, which is derived from the genus Strix (Olson and James, 1991). In the Strigiformes, as well as in the genus Accipiter, this lengthening of the legs corresponds to an adaptation for catching birds on islands lacking terrestrial mammals. This species must have had a very secretive life in the forests or remote areas, because the early explorers of Reunion never spoke of nocturnal raptors, although eared owls were noted historically on the other Mascarene Islands. Family STURNIDAE Genus Fregilupus Lesson Fregilupus varius (Boddaert, 1783) Reunion Starling MATERIAL.—Grotte des Premiers Francais: 1. d. femur, 1993-57. REMARKS.—This passerine femur agrees in size and morphological characteristics with what is known of the Reunion Starling. On the posterior face, the proximal edge of the internal condyle ends internally with a point that projects proximad. On the internal face, the internal condyle is anteroposteriorly compressed. This characteristic accentuates the shape of the external condyle, which looks very protmding, and the rotular groove on the anterior face is deep, as was indicated by Murie (1874), who described the only known skeleton of this species. The distal width (7.2 mm) and the distal depth (5.8 mm) agree with a femur the total length of which ranges from 31.6 mm (Berger, 1957) to 35.6 mm (Murie, 1874) or 37 mm (Gunther and Newton, 1879). The Reunion Starling became extinct between 1838 and 1858 (Barre and Barau, 1982). Relationships, Origin, and Fate of the Reunion Avifauna The avifauna found as fossils on Reunion differs from that of the other two Mascarenes (Table 13) in that, with the exception of Dryolimnas augusti, none of the species had lost their ability to fly. Lacking on Reunion are the most distinctive Mascarene birds, namely, the dodo and solitaire {Raphus, Pezophaps), the large flightless rails {Aphanapteryx, Erythromachus), and the large parrots with enormous bills and atrophied wings {Lophopsittacus mauritianus, Necropsittacus rodericanus) (Newton and Newton, 1876; Gunther and Newton, 1879). Among the forms that perhaps had lost the ability to fly is the Oiseau bleu, placed by Olson (1977) in the genus Porphyrio, and which was either an extinct species of that genus or a population of the modem species Porphyrio porphyrio (Linnaeus). Dubois (1674) said that it could not fly, but in 1724 Father Brown said that it was able to fly, but rarely and just above the ground (Barre and Barau, 1982, our translation). The authenticity of Father Brown's report has been questioned (Lougnon, 1970, 1992), but Cheke (1987) thinks that his report comes from an unidentified but authentic source. Remains referable to Oiseau bleu are yet to be found, so we know nothing more about it. The other genera represented on Mauritius and Rodrigues by species with reduced flying ability are represented on Reunion by species with normal flying ability. This is the case for Nycticorax, with the flying species N. duboisi, and for Falco duboisi, the coracoid and ulna of which are not reduced, unlike that of F. punctatus, of Mauritius. The other extinct species, namely

NUMBER 89 33 Family PHALACROCORACIDAE ARDEIDAE THRESKIORNITHIDAE PHOENICOPTERIDAE ANATIDAE ACCIPITR1DAE FALCONIDAE TURNICIDAE RALLIDAE RAPHIDAE COLUMBIDAE PSITTACIDAE STRJGIDAE APODIDAE HlRUNDINIDAE CAMPEPHAGIDAE PYCNONOTIDAE MUSCICAPIDAE TURDIDAE TlMALIIDAE SYLVIIDAE ZOSTEROPIDAE PLOCEIDAE STURNIDAE TABLE 13.—Native resident land birds of the Mascarene Islands. L=species still living on the island today, E=species completely extinct, X=species now extinct but known by modem specimens, F=species known from fossils found on the specified island, H=species known by historical accounts (when not known from fossils). (Egretta dimorpha on Mauritius is after Milne-Edwards (1874, pl. 33: fig. 3). There is no indication that "Necropsittacus" borbonicus belongs to the same genus as N. rodericanus. Hypsipetes (species undescribed) and Timaliinae (genus and species undescribed) on Rodrigues from Cowles (1987). A supposed grebe from Mauritius was based on a fossil of the migratory Whimbrel, Numenius phaeopus (Cowles, 1987).) Reunion Phalacrocorax africanus? (as "Cormoran"), H Nycticorax duboisi, E,F Egretta dimorpha? (as "Aigrette blanche et grise"), H Threskiornis solitarius, E,F Phoenicopterus ruber, F Alopochen (M.) kervazoi, E,F Anas theodori, E, F Aythya sp., F Circus maillardi, L Falco duboisi (as "Pieds jaunes"), E,F "Emerillons," E,H" Turnix nigricollis (as "Petites perdrix"), L,H Dryolimnas augusti, n. sp. E,F - Porphyrio caerulescens (as "Oiseau bleu"), E,H Fulica newtonii, E, F - Alectroenas sp.? (as "Pigeon couleur d'Ardoise"), E,H Nesoenas duboisi, E, F - Streptopelia picturata, L, F - "Perroquet gris," E,H "Necropsittacus" borbonicus (as "Perroquet vert a tete, dessus des ailes et queue couleur de feu"), E,H Mascarinus mascarinus, E,X,F Psittacula eques/echo? (as "Perroquet vert a collier noir"), H Mascarenotus grucheti, E,F Collocalia francica, L Phedina borbonica, L Coracina newtoni, L Hypsipetes borbonicus, L Terpsiphone bourbonnensis, L Saxicola tectes, L - - Zosterops borbonicus, L Zosterops olivaceus, L Foudia sp. (as "Moineaux"), E,H Fregilupus varius, E,X,F Threskiornis solitarius, Alopochen (M) kervazoi, Anas theodori, Nesoenas duboisi, and Mascarinus mascarinus, do not show reduction in the scapular girdle or wing bones. Alopochen (M) kervazoi shows a very slight reduction compared with the recent African forms, but this also exists in the extinct form from Madagascar. We suggest that this reduction had already occurred in the Malagasy forms before they colonized Mauritius Phalacrocorax africanus, F Nycticorax mauritianus, E, F Egretta dimorpha, F - Phoenicopterus ruber, F Alopochen mauritianus, E,F Anas theodori, E,F - Circus alphonsi, E,F Falco punctatus, L,F - - Dryolimnas cuvieri, X,F Aphanapteryx bonasia, E,F - Fulica newtonii, E, F Raphus cucullatus, E,X,F Alectroenas nitidissima, E,X,F Nesoenas mayeri, L, F - Streptopelia picturata, L, F, H Lophopsittacus mauritianus, E, F "Lophopsittacus" bensoni, E, F - Psittacula eques/echo, L, F Mascarenotus sauzieri, E,F Collocalia francica, L Phedina borbonica, L Coracina typica, L Hypsipetes olivaceus, L Terpsiphone bourbonnensis, L - — - Zosterops borbonicus, L Zosterops chloronothos, L Foudia rubra, L - Rodrigues - Nycticorax megacephalus, E,F - - possibly historical "geant," H - - - - - - - - Erythromachus leguati, E,F - - Pezophaps solitaria, E, F - - "Columba" rodericana, E,F Streptopelia picturata, L, F - - Necropsittacus rodericanus, E,F - Psittacula exsul, E,X,H Mascarenotus murivorus, E,F - - - Hypsipetes, sp. undescribed, E,F - TIMALIINAE, genus and sp. undescribed, E,F Acrocephalus rodericanus, L - - Foudia flavicans, L Necropsar rodericanus E,F Reunion. Fulica newtonii shows a reduction of the coracoid and wing bones, but it is the same species as in Mauritius, and it probably colonized Reunion from that island. Mascarenotus grucheti has a more reduced humems than does M. sauzieri, but it is the only exception. Reunion Island dates back 3 Ma (Molnar and Stock, 1987), a length of time amply sufficient for birds to lose their flying

32 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY<br />

Without any justification, Rothschild (1907) placed this form<br />

in the genus Necropsittacus of Rodrigues and described it as N.<br />

borbonicus.<br />

Family STRIGIDAE<br />

Genus Mascarenotus Mourer-Chauvire, Bour,<br />

Moutou, and Ribes<br />

Mascarenotus grucheti Mourer-Chauvire, Bour,<br />

Moutou, and Ribes, 1994<br />

Gmchet's Mascarene Owl<br />

FIGURE 13^<br />

MATERIAL.—Grotte des Premiers Francais. Holotype: r.<br />

tarsometatarsus, 1993-49. Paratypes: 1. humems, 1993-50; r. p.<br />

tibiotarsus, 1993-48; r. d. tibiotarsus, 1993-47.<br />

Grotte de l'Autel: r. tarsometatarsus, 330537; r. d. tarsometatarsus,<br />

330538.<br />

Grotte "au sable": 1. quadrate, 330809; 1. p. femur, 330737.<br />

Marais de l'Ermitage: r. tarsometatarsus, 1800.<br />

REMARKS.—We have not found any additional remains other<br />

than those used in the original description of Mascarenotus<br />

grucheti (Mourer-Chauvire et al., 1994). We have placed all<br />

the Mascarene strigiforms in the extinct genus Mascarenotus,<br />

which resembles the genus Otus but presents some distinctive<br />

morphological characteristics. This genus includes one species<br />

on Mauritius, Mascarenotus sauzieri, based on Strix sauzieri<br />

Newton and Gadow, 1893 (synonyms, Otus commersoni Oustalet<br />

(1896), Strix newtoni Rothschild (1907)), one species on<br />

Rodrigues, Mascarenotus murivorus based on Strix {Athene)<br />

murivora Milne-Edwards (1874) (synonym, Bubo (?) leguati<br />

Rothschild (1907)), and one species on Reunion, Mascarenotus<br />

grucheti Mourer-Chauvire et al., 1994.<br />

The tarsometatarsi of M. grucheti are very close in size to<br />

those of M. sauzieri, although the only humems known from<br />

Reunion is clearly smaller than those of M. sauzieri, for which<br />

we have a good sample of comparative material (12 specimens).<br />

The ratio of element lengths (Mourer-Chauvire et al.,<br />

1994, fig. 1) shows M. sauzieri to be strikingly parallel to modem<br />

insular species of the genus Otus, such as Otus lawrencii<br />

(Sclater and Salvin), or O nudipes (Daudin), from the West Indies,<br />

in which the legs are much more elongated than in continental<br />

forms. The same adaptation is found in the four extinct<br />

insular species of the genus Grallistrix, from Hawaii, which is<br />

derived from the genus Strix (Olson and James, 1991). In the<br />

Strigiformes, as well as in the genus Accipiter, this lengthening<br />

of the legs corresponds to an adaptation for catching birds on<br />

islands lacking terrestrial mammals.<br />

This species must have had a very secretive life in the forests<br />

or remote areas, because the early explorers of Reunion never<br />

spoke of nocturnal raptors, although eared owls were noted historically<br />

on the other Mascarene Islands.<br />

Family STURNIDAE<br />

Genus Fregilupus Lesson<br />

Fregilupus varius (Boddaert, 1783)<br />

Reunion Starling<br />

MATERIAL.—Grotte des Premiers Francais: 1. d. femur,<br />

1993-57.<br />

REMARKS.—This passerine femur agrees in size and morphological<br />

characteristics with what is known of the Reunion<br />

Starling. On the posterior face, the proximal edge of the internal<br />

condyle ends internally with a point that projects proximad.<br />

On the internal face, the internal condyle is anteroposteriorly<br />

compressed. This characteristic accentuates the shape of the<br />

external condyle, which looks very protmding, and the rotular<br />

groove on the anterior face is deep, as was indicated by Murie<br />

(1874), who described the only known skeleton of this species.<br />

The distal width (7.2 mm) and the distal depth (5.8 mm) agree<br />

with a femur the total length of which ranges from 31.6 mm<br />

(Berger, 1957) to 35.6 mm (Murie, 1874) or 37 mm (Gunther<br />

and Newton, 1879).<br />

The Reunion Starling became extinct between 1838 and<br />

1858 (Barre and Barau, 1982).<br />

Relationships, Origin, and Fate of the Reunion Avifauna<br />

The avifauna found as fossils on Reunion differs from that of<br />

the other two Mascarenes (Table 13) in that, with the exception<br />

of Dryolimnas augusti, none of the species had lost their ability<br />

to fly.<br />

Lacking on Reunion are the most distinctive Mascarene<br />

birds, namely, the dodo and solitaire {Raphus, Pezophaps), the<br />

large flightless rails {Aphanapteryx, Erythromachus), and the<br />

large parrots with enormous bills and atrophied wings {<strong>Lo</strong>phopsittacus<br />

mauritianus, Necropsittacus rodericanus) (Newton<br />

and Newton, 1876; Gunther and Newton, 1879). Among<br />

the forms that perhaps had lost the ability to fly is the Oiseau<br />

bleu, placed by Olson (1977) in the genus Porphyrio, and<br />

which was either an extinct species of that genus or a population<br />

of the modem species Porphyrio porphyrio (Linnaeus).<br />

Dubois (1674) said that it could not fly, but in 1724 Father<br />

Brown said that it was able to fly, but rarely and just above the<br />

ground (Barre and Barau, 1982, our translation). The authenticity<br />

of Father Brown's report has been questioned (<strong>Lo</strong>ugnon,<br />

1970, 1992), but Cheke (1987) thinks that his report comes<br />

from an unidentified but authentic source. Remains referable to<br />

Oiseau bleu are yet to be found, so we know nothing more<br />

about it.<br />

The other genera represented on Mauritius and Rodrigues by<br />

species with reduced flying ability are represented on Reunion<br />

by species with normal flying ability. This is the case for Nycticorax,<br />

with the flying species N. duboisi, and for Falco duboisi,<br />

the coracoid and ulna of which are not reduced, unlike that<br />

of F. punctatus, of Mauritius. The other extinct species, namely

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