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NUMBER 89<br />

Puffinus Iherminieri Lesson, 1840<br />

Audubon's Shearwater<br />

MATERIAL.—Grotte des Premiers Francais: Very numerous<br />

remains from all parts of the skeleton, corresponding at<br />

least to 1 adult in layer 4 and to 27 individuals (14 adults, 13<br />

juveniles) in layer 5.<br />

Grotte "au sable": Very numerous remains, corresponding<br />

at least to 11 individuals (2 adults, 9 juveniles):<br />

330572-330588, 330591, 330596-330597, 330599,<br />

330628-330638, 330642-330643, 330645-330646,<br />

330651-330652, 330661-330664, 330668, 330670,<br />

330684-330699, 330712-330726.<br />

Grotte de l'Autel and Grotte "au sable": Very numerous remains,<br />

corresponding at least to 4 individuals (2 adults, 2 juveniles):<br />

330745-330762.<br />

REMARKS.—The high proportion of juveniles indicates that<br />

both species of Puffinus were probably nesting in these cavities<br />

or their surroundings. Puffinus pacificus now nests at Reunion<br />

only on a small islet, accessible only with difficulty for humans<br />

or introduced mammals, whereas P. Iherminieri nests in numerous<br />

parts of the island (Jouanin, 1987). In the three caves,<br />

P. pacificus is more numerous than P. Iherminieri, but at<br />

present P. Iherminieri is much more common on the island.<br />

Two other species of Procellariidae, Pseudobulweria aterrima<br />

(Bonaparte) and Pterodroma baraui (Jouanin), now nest on<br />

Reunion, at high altitudes. Subfossil remains of P. baraui have<br />

been found in the Caverne a Cotte, at about 1800 m elevation<br />

(Jouanin and Gill, 1967). In our fossil localities, which are situated<br />

at low elevation, we did not find remains that can be attributed<br />

to either of these species. The limited fossil evidence<br />

therefore suggests that they may always have nested at higher<br />

elevations than Puffinus.<br />

Family PHAETHONTIDAE<br />

Genus Phaethon Linnaeus<br />

Phaethon lepturus Daudin, 1802<br />

White-tailed Tropicbird<br />

FIGURE 4/<br />

MATERIAL.—Grotte des Premiers Francais: Part of skull,<br />

1993-51.<br />

Grotte de l'Autel: 1. coracoid, 330515 (Figure At).<br />

REMARKS.—The internal length of the coracoid is 39.0 mm,<br />

which falls within the range of variation of modern P. lepturus<br />

(36.2^10.1, «=10) and is clearly smaller than in the other two<br />

species, P. rubricauda Boddaert, the Red-tailed Tropicbird<br />

(47.0-53.9, /i=24), and P. aethereus Linnaeus, the Red-billed<br />

Tropicbird (47.0-50.4, «=7). Phaethon lepturus still nests on<br />

Reunion.<br />

Family ARDEIDAE<br />

Genus Nycticorax Forster<br />

Megaphoyx Hachisuka 1937b: 148 [type by original designation, Ardea megacephala<br />

Milne-Edwards, 1874].<br />

Each of the three Mascarene islands sustained an extinct species<br />

of Nycticorax. The first of these species was described<br />

from Rodrigues by Milne-Edwards (1874) under the name<br />

Ardea megacephala. Giinther and Newton (1879) showed that<br />

the Rodrigues heron belonged in the genus Nycticorax and that,<br />

although its size was not very different from a large modern N.<br />

nycticorax (Linnaeus), its wing bones were proportionally<br />

shorter and its femur, tarsometatarsus, and pedal phalanges<br />

proportionally longer. Actually, when compared with typical<br />

N. nycticorax nycticorax, the wings are not very short, but the<br />

femora, tibiotarsi, and tarsometatarsi are wider, longer, and<br />

more robust (Cowles, 1987). Newton and Gadow (1893) described<br />

a second, Mauritian species, Butorides mauritianus,<br />

which is smaller than N. megacephalus, and then Rothschild<br />

(1907) described a third species, from Reunion, under the name<br />

of Ardea duboisi. Rothschild placed the three species in the genus<br />

Ardea but wrote (1907:115): "From the short, stout legs<br />

and general build, I am inclined to think that all three of these<br />

Herons belong to the genus Nycticorax." Later, other authors<br />

(Lambrecht, 1933; Hachisuka, 1953; Brodkorb, 1963) placed<br />

the three species in different genera, and it was Cowles (1987)<br />

who first formally united them in the genus Nycticorax.<br />

Nycticorax duboisi (Rothschild, 1907), new combination<br />

Reunion Night Heron<br />

FIGURE 4a-h<br />

"Butors ou Grands Gauziers" Dubois, 1674:169.<br />

Ardea duboisi Rothschild, 1907:114 [based on birds described by Dubois<br />

(1674) from Bourbon (= Reunion Island)].<br />

Megaphoyx duboisi.—Hachisuka, 1953:175.<br />

Nycticorax n. sp. Cowles, 1987:94.<br />

Nycticorax borbonensis Cowles, 1994:90, fig.ld,e [new synonymy; holotype,<br />

distal half of left tibiotarsus MNHN, LAC 1993-35, from bed 4, Grotte des<br />

Premiers Francais (Grande Caverne), Reunion Island].<br />

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

borbonensis (see below).<br />

Grotte de l'Autel: r. scapula, 330516.<br />

Marais de l'Ermitage: r. scapula, 1866; 1. p. coracoid,<br />

1831; 1. d. humerus, 1826; r. ulna, 1828; 1. p. ulna, 1832; r. femur,<br />

1827; 2 r. d. tibiotarsi, 1829, 1830; r. tarsometatarsus,<br />

1916; 2 1. d. tarsometatarsi, 1865, 1917.<br />

DESCRIPTION AND COMPARISONS.—The material from<br />

Reunion agrees perfectly with the genus Nycticorax. The different<br />

species of this genus show much variation in size, the<br />

largest being N. caledonicus (Gmelin), which lives in Indonesia,<br />

Australia, New Zealand, New Caledonia, and in some Pacific<br />

archipelagos (Mayr and Cottrell, 1979). The remains of N.

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