Er Ve JE Volume 35, 1965 N° Spécial \ W L’OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE LE PROFESSEUR BERLIOZ Volume Jubilaire REVUE TRIMESTRIELLE DE LA SOCIÉTÉ ORNITHOLOGIQUE DE FRANCE Rédaction : 55, rue de Buffon, Paris (Ve) Source : MNHN. Paris L'OISEAU ET LA REVUE FRANCAISE D’ORNITHOLOGIE Publié avec le concours du G.S.C. et de l'O.R.S.T.O.M. Comité de lecture : MM. J. BERLIOZ, R.-D. ETCHECOPAR et M. LEGENDRE Abonnement annuel : France et Etranger : 34 F. Toute correspondance concernant la Revue doit être adressée au Secrétariat : 55, rue de Buffon, Paris (V°). Tout envoi d’argent doit être adressé au nom de la « Société Ornithologique de France ». Compte Chèques Postaux Paris 544-78. AVIS IMPORTANT Par suite de la récupération d’un certain stock, nous avons le plaisir d'annoncer à nos membres que nous pou- vons mettre à leur disposition les années arriérées suivantes de « L'Oiseau et la R.F.0. », au prix de : 45,00 F - 1935 - 1937 - 1946 - 1954 - 1956. 30,00 F - 1933 - 1938 - 1949 - 1957. 20,00 F - 1955 - 1958 - 1959 - 1960 - 1961 - 1962. 12,00 F - 1942 - 1947 - 1951. La rédaction ne prend sous sa responsabilité aucune des opi- nions émises par les auteurs des articles insérés dans la Revue. La reproduction, sans indication de source ni de nom d’auteur, des articles publiés dans la Revue est interdite. Les auteurs sont priés d'envoyer leurs manuscrits dactylogra- phiés, sans aucune indication typographique. Source : MNHN. Paris Volume 35, 1965 N° Spécial L’OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE LE PROFESSEUR BERLIOZ Volume Jubilaire BIBL.DU MUSEUM D'HIST, LATUR, LD Ÿ REVUE TRIMESTRIELLE DE LA SOCIÉTÉ ORNITHOLOGIQUE DE FRANCE Rédaction : 55, rue de Buffon, Paris (Ve) Source : MNHN. Paris L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE SOMMAIRE AVANT-PROPOS . . à Ë J. Dorsr : Le Professeur Berlioz 4 4 4 #40 ni à ne © 3 D. AMADON : Notes on the Galapagos Hawk . . + + + . + : . 9 D. BANNERMAN : Some reflections on two recent visits to the Azores archi- pelage “078 20h eanieaifr 5 olensianr oieduiit 22 G. J. BROEKHUYSEN : An analysis of bird casualties on the roads in the South Western Cape Province, South Africa . . . . . . 35 J. DELAGOUR : Les Pigeons de la Nouvelle-Calédonie . . . + + - : 52 G. P. Demenriry et L. S. STÉPANYAN : Quelques particularités de la reproduction des oiseaux dans les hautes zones du Thian-Chan . . : . : . 58 A. GHiGt : Sur le Faisan de Hume et le genre Calophasis 7; 65 S. HüRSTADIUS : De l'effet désastreux de l’enrobage des semences sur l'avi- faune AMÉOISE Lee md dar an le 71 D. Lack and R. E. MOREAU : Clutch-size in tropical passerine birds of forest and 2 Li Source : MNHN. Paris E. Mayr : Classification, identification and sequence of genera and De CS DR ie ee NU | G. NIETHAMMER : Zur Taxonomie von Cinclus cinclus. Ein tiergeogra- phisthés Problem Et ei PAR dt nr 96 S. Dillon RiPLEY : Le Martinet pâle de Socotra . . . . . . . . . . 101 F. SALOMONSEN : Notes on the Sunbird-Asitys (Neodrepanis) . . . . . 103 C. G. SIBLEY : Molecular systematics : New techniques applied to old problems HSE, or E. und V. STRESEMANN : Die Mauser des Kampfläufers Philomachus pugnaæ . . . 125 A. Landsborough THOMSON : The transequatorial migration of the Manx Shearwater (Putindes Anglais) + — =. … Merci... 100 G. VAURIE : Distribution régionale et altitudinale des genres Garrulax et Babax et notes sur leur systématique . . . . . 141 K. H. Voous : Specimens of Lear’s Macaw in the Zoological Museum of ECC RE DE TR EN SR EN Nu A. WETMORE : Additions to the list of birds of the Republic of Colombia. 156 J. M. WiINTERBOTTOM : A review of the southern subspecies of the Fawn-coloured Lark Mirafra africanoides Smith . . . . . . . . 163 Source : MNHN. Paris ET LA © FRANÇAISE D'ORNITHOLOGIE Source : MNHN., Paris AVANT-PROPOS Depuis longtemps déjà la Société Ornithologique de France désirait manifester sa reconnaissance au Professeur BERLIOZ pour l’appui que celui-ci n’a cessé de lui prodiguer au cours de sa longue carrière au Muséum National d'Histoire Natu- relle. Sa retraite officielle apporte aujourd’hui l’occasion recherchée. Notre Conseil d'Administration avait d’abord envisagé de faire frapper une médaille à l'effigie de celui qui fut pendant si longtemps le titulaire de la seule chaire d’ornithologie fran- çaise ; mais sa discrétion et sa modestie nous a fait penser que nous avions plus de chance de lui être agréable si nous lui dédions un fascicule de cette revue dont il dirigea si heu- reusement la rédaction pendant plus de vingt années. Nous savions que dans ce but il serait aisé d'obtenir la contribution des plus grands noms de l’ornithologie contem- poraine, mais des impératifs matériels nous obligèrent à res- ter dans des limites hélas bien modestes. C’est la raison pour laquelle nous ne nous sommes adressés qu'aux seuls Prési- dents et Secrétaires Généraux des Congrès internationaux d’ornithologie. On se rappelle en effet que le Professeur Ber- L10Z présida lui-même le 12° Congrès, qui tint ses assises à Helsinki en 1958. A ces personnalités nous avons ajouté les ornithologistes des grands musées mondiaux, avec lesquels le Professeur BERLIOZ maintenait un contact permanent. Les réponses favorables à notre appel furent si nom- breuses et si sincères que nous fûmes amenés très vite à réa- liser notre projet. Nous tenons à remercier ici au nom de tous nos membres ceux qui ont accepté de participer à cet hom- Source : MNHN. Paris 2 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE mage envers l’un des leurs. Il faut voir dans ce généreux élan la preuve de l'estime générale dont jouit notre éminent col- lègue. Elle nous paraît d'autant plus justifiée que son atta- chante personnalité sait joindre à ses qualités professionnelles celles du cœur et de l'esprit. Soubaitons que son amour des voyages d’étude et ses goûts d'indépendance ne l’éloignent pas trop souvent de nous, et qu’il puisse ainsi continuer à faire profiter notre Société de sa grande expérience et de ses profondes connaissances dans un domaine scientifique qui est le nôtre. Le Président : Le Secrétaire Général : F. BOURLIÈRE. R. D. ErcHÉCOPAR. Source : MNHN. Paris LE PROFESSEUR BERLIOZ par Jean DorsT Muséum National d'Histoire Naturelle (Paris) En même temps qu'un grand honneur, c’est un devoir bien difficile que de présenter ce recueil d'articles rédigés par quelques-uns des représentants les plus qualifiés de l'ornitho- logie mondiale et publiés par notre Revue en hommage au Professeur BErLioZ. Pour le Comité de Rédaction, ce fut une tâche bien plus difficile encore que d'établir la liste des per- sonnalités à pressentir, car nombreux sont ceux qui auraient voulu y collaborer. Il a fallu notamment renoncer à la par- ticipation des ornithologistes français. Je le déplore à beau- coup de points de vue, car nombreux sont ceux qui auraient voulu apporter leur hommage à celui qui fut leur maître et auquel on doit le renouveau d'activité ornithologique au Muséum. En définitive n’ont été sollicités que quelques-uns des ornithologistes étrangers qui occupèrent des fonctions dans les Congrès internationaux en tant que Présidents ou Secré- taires généraux. Ce n’est pas ici la place d’énumérer les mérites du Profes- seur BerLioz. Je voudrais simplement souligner ce que lui doit le laboratoire d’Ornithologie du Muséum. Fréquentant notre établissement depuis 1905, il y est entré comme jeune assistant en 1920. Il trouva l’ornithologie dans un état de sommeil, car le titulaire de la chaire, mammalogisle de spé- cialité, ne lui avait pas donné l'impulsion nécessaire. Dès 1926, le Professeur Bourpezze, dont il fut le collaborateur direct pendant plus de 20 ans, lui abandonna immédiatement l’en- tière responsabilité du département d’ornithologie. La tâche était énorme. Les collections, qui rivalisaient avec celles des établissements étrangers similaires du temps de Miine- Epwanrps, n'avaient guère été augmentées depuis et n'étaient L'Oisean et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris 4 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE pas rangées comme elles auraient dû l’être. Pendant ce temps les grands musées d’outre-Atlantique avaient pris une avance qui rendait la comparaison peu flatteuse pour la France. Patiemment M. BERLIOZ commença la détermination, le classement et le rangement méthodique des spécimens jusque- là entreposés sans grand ordre. Son attention se porta simul- tanément sur la collection publique, qu’il s’astreignit à rendre plus attrayante. Il en extraya les types en mettant ces « éta- lons » des espèces à l'abri des méfaits d’une exposition à la lumière et à la poussière. Ces collections anciennes du Muséum renfermaient également bon nombre de spécimens représen- tant des espèces actuellement disparues ou en voie d’extinc- tion. Ses efforts permirent de disposer dans une salle parti- culière ce matériel précieux à l’occasion du Tricentenaire du Muséum en 1935. Un de ses grands regrets fut de ne pouvoir mener à bien une complète transformation des galeries publiques, faute de crédits et aussi du fait d’une certaine désaffection des autorités responsables. Ce travail reste à entreprendre, car c’est un devoir pour notre Maison que d'of- frir au grand public des collections attrayantes, esthétiques et didactiques, moyen efficace d'enseignement populaire. La position du Muséum, encadré par les nouvelles Facultés des Sciences et des Lettres, augmente aussi nos responsabilités vis-à-vis des étudiants. Mais une collection scientifique est morte si elle ne s’ac- croît pas continuellement. De grandes lacunes restaient à com- bler pour faire de nos collections nationales un outil de tra- vail et une source de documentation universelle. M. BERLIOZ s’attacha à cet enrichissement. Une partie des apports nou- veaux provint de quelques grands voyageurs et chargés de ion, avant tout de M. Jean DELACOUR avec qui se noua une collaboration très suivie. Une véritable campagne de pros- pection fut aussi entreprise parmi les collecteurs profession- nels à travers le monde. De longues séries d'oiseaux affluèrent au Muséum, en provenance notamment d’Ecuador, du Mexique et de quelques pays africains. Des échanges réalisés avec d’autres élablissements élargirent encore cet échantillon- nage, pour faire des collections nationales le centre de docu- mentation moderne et complet dont rêve tout conservateur. Le Professeur BERLIOZ est cependant bien autre chose qu’un savant de cabinet. Naturaliste dans l’âme, il sait depuis toujours que c’est par le contact direct avec les êtres vivants Source : MNHN. Paris LE PROFESSEUR BERLIOZ 5 qu'on a le plus de chances d'apprendre la véritable histoire naturelle. Dès 1924, il entreprit de longs voyages à travers le monde et, à part l'Océanie, il est peu de régions qu’il nait parcourues. L'Amérique du Sud, puis surtout l’Asie, retinrent plus particulièrement son attention, cette dernière partie du monde ayant pour lui le charme supplémentaire de peuples de haute culture et de civilisations millénaires. Il recueillit ainsi au fil des années une documentation considérable sur la répartition et les mœurs des oiseaux, des autres animaux aussi, en même temps qu'il nouait des contacts fructueux pour notre Maison avec des correspondants répartis dans le monde entier. Cette double orientation, motivée par deux tendances com- plémentaires, détermina ses recherches personnelles. Collec- tionneur dans l'âme, il est avant tout un systématicien, car il sait que, si la systématique est le début de toute science naturelle, elle en est aussi la fin. Beaucoup de ses travaux —— en tout plusieurs centaines — ont porté sur la taxinomie ornithologique. Il a décrit de nombreuses formes nouvelles. Et si les Trochilidés ont retenu tout spécialement son atten- tion, l’ensemble des oiseaux n’a pas échappé à ses investiga- tions, avec toutefois une prédilection affirmée pour ceux des régions tropicales, encore relativement mal connus. Un tableau d'ensemble de la systématique ornithologique fut dressé sous sa signature dans le Traité de Zoologie du Profes- seur GRASSÉ. Une autre partie de ses travaux porte sur la biogéographie, science vers laquelle le portaient ses goûts et ses connais- sances de terrain. Après avoir étudié une avifaune, il se plaît à en étudier la composition, à préciser ses affinités et à démêler la manière dont elle s’est formée, sans jamais se lancer dans les hypothèses hasardeuses de « science-fiction » où se plaisent beaucoup de biogéographes. Le réel a toujours plus d’attrait pour lui que la projection dans le passé ou l'avenir. Les voyages du Professeur BERLIOZ lui avaient aussi appris à quel point la nature était menacée dans le monde et partout il avait constaté avec anxiété la régression de la faune et de la végétation devant l'avance de la civilisation technique. Ses missions au Canada et aux Etats-Unis lui avaient montré, dès 1924, combien ces pays étaient en avance sur nous dans le domaine de la protection des espèces et des habitats sauvages. Source : MNHN. Paris 6 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Aussi devint-il un ardent apôtre de ces idées, nouvelles à cette époque en France. S'il fut un des premiers à les diffuser dans notre pays, son intérêt pour ces questions n’a jamais faibli, comme le démontre une participation active et suivie à tous les mouvements de défense de la nature. Mais le Professeur BERLIOZ est plus que cela encore. C’est un naturaliste complet, formé à la Faculté de Pharmacie et à celle des Sciences, où il s’est imprégné de toutes les disci- plines dont le large faisceau forme les sciences naturelles. Il a continué à s'intéresser à beaucoup d’entre elles et ses con- naissances en botanique et surtout en entomologie ne le cèdent guère à celles qui lui sont nécessaires pour sa vie profession- nelle. Ce qui est beau l’attire plus que le reste, et ce n’est pas en vain que ses préférences vont aux Oïiseaux-mouches, aux Cétoines, aux Papilio et à quelques minéraux. Mais les autres groupes ne l’intéressent guère moins et nul sans doute plus que lui n’a ressenti l'unité du monde vivant et la néces- sité de ne pas en dissocier une partie si l’on veut comprendre les phénomènes de la vie et la place de chaque être dans la communauté. Les charges du Professeur BERLIOZ ont été lourdes. Pen- dant de longues années les crédits furent mesurés ; l’expan- sion scientifique, lente dans l’ensemble en France, ne touchait guère les disciplines fondamentales jugées archaïques. Il eut donc à lutter pour accroître les collections et dégager les cré- dits nécessaires à leur présentation. Quand il prit la direction de la chaire, nous sortions de la période difficile de la guerre et les années qui suivirent les arrêts des hostilités ne le furent guère moins. Ici encore il s’agit de trouver les fonds nécessaires à la reprise des acti- vités normales du laboratoire et à son développement. IL s’y employa sans faire de bruit et en dépit de multiples écueils développa le laboratoire en donnant à ses collaborateurs les possibilités de travail qui avaient fait défaut à leurs prédé- cesseurs. Le renom des travaux du Professeur BERLIOZ, les contacts qu’il avait pris à travers le monde, lui valurent de nombreux honneurs qu’il n’avait jamais recherchés. Membre d'honneur de toutes les grandes sociétés ornithologiques du monde, il fut appelé en 1958 à présider le XII° Congrès international d’Ornithologie réuni à Helsinki. L'adresse présidentielle qu’il adressa à cette occasion reste présente à l'esprit de tous, car Source : MNHN. Paris LE PROFESSEUR BERLIOZ 7 elle synthétise un aspect important de sa pensée : le rôle des collections dans l’avenir de l’ornithologie. Il insista sur le rôle des spécimens de collection comme base la plus solide de nos connaissances relatives au peuplement animal et à son évolu- tion actuelle. Sans mettre en doute la valeur des études de terrain et de la biologie expérimentale, il affirma que celles- ei sont vaines si elles ne reposent pas sur les solides con- naissances systématiques que seules peuvent donner des col- lections. Cet appel résonna longtemps à une époque où l’on glissait vers des routes sans issues. Le renouveau actuel de la systématique ornithologique, en particulier aux Etats-Unis, lui donne entièrement raison. Le Professeur BERLIOZ a aussi su depuis toujours qu'un chercheur reste sans grande action s’il ne sait pas s’entourer d'élèves et de collaborateurs formés à son école. Il a su sus- citer des enthousiasmes autour de lui. Les jeunes n’abordent le Muséum qu'avec un respect qui touche à la crainte tant Vatmosphère leur paraît chargée de science et d’austérité. Il a su leur parler, les mettre à leur aise, les placer en face des problèmes et leur confier des collections sous sa garde rigou- reuse, pour apprendre et étudier. Il n’a pas craint de perdre un temps précieux que d’autres maîtres auraient consacré à la recherche ou à courir après de vains honneurs. Il les a formés à la dure école du laboratoire, et a toujours compris que ses élèves s’échappent sur le terrain, munis de la forma- tion solide qui ne s’acquiert que par la manipulation de spé- cimens. En dépit d'opinions solidement établies, il a toujours admis la contradiction et les longues discussions, souvent animées, qui eurent le laboratoire pour théâtre, apportèrent beaucoup à ses interlocuteurs. Les hommes de science sont souvent personnels, exclusifs, voire égoïstes. Le Professeur BERLIOZ est exactement le contraire et ses jeunes élèves — ou ceux qui furent jeunes — lui savent gré d'être un maître compréhensif, sensible, parfois taquin, en même temps que ferme et érudit. C'est en témoignage d'estime, d’admiration et de respec- tueuse affection que les pages de ce fascicule ont été écrites. Elles ne marquent pas la fin d’une carrière. Car un natura- liste du Muséum n’est jamais à la retraite. Il arrive un âge où la société le décharge de ses obligations administratives, un devoir combien pesant pour un homme dont la vocation fut déterminée par la recherche et non par la rédaction de Source : MNHN. Paris 8 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE rapports et de demandes de crédits, de préparation de budgets et de bordereaux de service. Un naturaliste continue toute sa vie le même travail, pour sa satisfaction personnelle sans doute, mais surtout pour connaître et faire connaître aux autres quelques-uns des secrets de la nature. Nul ne le fait mieux que le Professeur BERLIOZ. Il y a quelques jours je le voyais étudier avec passion une collection fraîchement arrivée du Pérou. Il est certain que son enthousiasme était le même que celui dont il témoignait quand, jeune néophyte, il pénétra dans la vieille maison de la rue de Buffon qu'il a su rénover dans le respect des traditions. Le Professeur BERLIOZ a su faire œuvre de naturaliste et de chef d’école. Faisant alterner travail en laboratoire et longs voyages à travers le monde, il a su mener une vie heureuse et calme, en même lemps que fructueuse et stimulante pour ceux qui l'ont entouré, un cas rare dans notre monde moderne. Ce recueil de travaux ne fait que marquer une étape de sa carrière. Au nom de la Société Ornithologique de France, au nom de tous ses collègues, de ses amis et de ses élèves, je le prie de bien vouloir l’accepter comme hommage de respect et d’affection. Source : MNHN. Paris L'OI ET LA FRANÇAISE D'ORNITHOLOGIE PL. I U REVU à Somerset West, Le Professeur Bentroz photographiant des Prot Province du Cap, Afrique du Sud, en septembre 1956 (Photographie G. J. Broekhuysen) Source : MNHN. Paris NOTES ON THE GALAPAGOS HAWK by Dean AMADON American Museum of Natural History, New York (Contribution n° 40 de la Fondation Charles Darwin pour les Galapagos) During field work in the Galapagos Islands in January and February 1964 while a member of the Galapagos Inter- national Scientific Project sponsored by the National Science Foundation, Grant Number GE 2370, and by the University of California, Berkeley, special attention was given to Buteo galapagoensis. Observations were carried out on this raptor at Punta Espinosa on Fernandina (Narborough) Island, January 26-February 5; above Tagus Cove, Isabela (Albe- marle) Island on January 30th ; on Española (Hood) Island from February 15-20 and finally for a couple of hours on Santa Fe (Barrington) Island on February 20. During two weeks or so on Santa Cruz (Indefatigable) this hawk, which is now rare, at least in the settled portions of that island, was not seen. Robert Pyle of our party did see one in the highlands. RELATIONSHIP OF THIS SPECIES A special interest in the systematics of hawks may serve as an excuse for commenting on this question at some length. Ripaway (1875) divided the genus Buleo into two subgenera, separated primarily by whether three, or four, outer prima- ries are notched. Buteo galapagoensis has three primaries notched and was placed in a subgenus Craxirex Gould, an exclusively American group, whose other species are platyp- terus, swainsoni, albicaudatus, polyosomus and (if a species) poecilochrous. The last four species named, along with gala- pagoensis constitute, 1 agree with RipGway, a natural unit, L'Oiseau et R.F.0., V. 85, 1965, n° spécial. Source : MNHN. Paris 10 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE indeed they may be a superspecies. Buleo swainsoni, is much weaker footed than the others, and it is probably a mistake to regard it as the closest relative of galapagoensis. RipGway placed all the other species of Buleo, including all the Old World ones, in the nominate subgenus, characte- rized by four notched primaries. It may be remarked, parenthetically, that, as a practical matter, the use of this character is not always as easy as it sounds, particularly in museum skins. Many specimens have the outer primaries in molt; the notch on the feathers becomes less pronounced in worn plumage, and some indi- viduals and, I suspect, even species may be intermediate, To illustrate these difficulties Rineway placed Buteo brachyurus in the group with four notched primaries ; the few I have examined seem to have only three, but perhaps I have yet to see one in first class condition. I should also add that Rincway (1873) recognized a genus Rupornis (five primaries notched) with species magnirostris and leucorrhous and a genus Asturina (four primaries not- ched) for the species nifidus. These three species are now generally, and I believe correctly, included in Buteo, but they admittedly (especially nitidus and magnirostris) are somewhat intermediate lowards some of the seemingly pri- mitive buteonine genera, notably Leucopternis and Buteogal- lus. If the above remarks are correct, a phylogenetic arrange- ment of the species of Buteo would begin with the three spe- cies just mentioned, then the large four-notched group, and finally the three-notched « Craxireæ » group. This does not seem very satisfactory. The four-notched group, ineluding all the Old World species and such New World ones as jamaïi- censis and regalis, seems to represent {he more advanced segment of the genus. This group seems to be the most like the related, more advanced « booted » eagles of such genera as Spizaetus, Hieraaetus and Aquila. One is led to suggest that the number of notched primaries may not be a very good clue to relationship. An examination of all the buteonine and aquiline genera of the family will, I believe, convince anyone that such is the case, though I shall not tabulate extensive evidence. This is not to say that the character is of no use ; it is one of the features showing the Craxirex group to be a monophylelic unit. It will also, of course, remain a useful Source : MNHN. Paris NOTES ON THE GALAPAGOS HAWK 11 aid in identifying some puzzling specimens, for exemple, cer- tain immatures of swainsoni. JOHNSON and PEETERS (1963) have recently set up a group of (American) woodland buteos, with the species magniros- tris, ridgwayi, lineatus platypterus and nitidus. They do not mention the question of primary-notching but obviously con- sider it of no importance, since their group contains species of each type, e. g., lineatus (4), and platypterus (3). As already intimated, 1 agree that this character is probably not always significant, but it has entered into discussions of the genus so extensively that it can hardly be ignored. JOHNSON and PEETERS have not mentioned either Buteo leucorrhous or B. brachyurus (and albigula, if specifically dis- tinct from brachyurus). The former I would think surely and the latter probably should be associated with their group. In the overall arrangement of the genus Buteo 1 would begin with this « woodland » group with the additions noted ; follow it by the « Craxirex » group, which may be regarded as a natural group of fairly primitive buteos which happen to have three primaries notched, and finally come the remainder of the genus beginning with such slightly aberrant species as albonotatus and solitarius and conclu- ding with the Old World and African ones. So far as the sequence of the « woodland » group is con- cerned, it should begin, not end, with the aberrant nitidus, which, as noted above, provides a link with Leucoplernis and, in the immature, Buteogallus. Some may argue that RipGway was correct and that the similarity of lineatus and platypterus, for example, is the result of ecological factors. While this may be true, we do have the fact that they are very similar and live in the same part of the globe. I would thus suggest the following sequence of species for the genus Buleo : nitidus, magnirostris, leucorrhous, ridgwayi, linea- lus, platypterus, brachyurus, swainsoni, galapagoensis, albi- caudatus, etc. Hagirs After reading of the tameness of the Galapagos Hawk and of how it eats centipedes and carrion, I had expected a sluggish, inoffensive bird. During the breeding season, at least, it is aclually a vigorous, noisy species and, as will be Source : MNHN. Paris 12 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE seen from the section on food below, it catches some large prey. In these respects it is to be compared with such a spe- cies as the Red-tailed Hawk, Buteo jamaicensis. In physical appearance also, the Galapagos Hawk is a more impressive bird than I, at least, had expected from exa- mination of frowzy museum specimens, often in immature plumage. As is now known, the adult plumage is always black ; phases do not occur. In the hand it is a rather brownish black, but this is not apparent in he field. In a pair under observation in the field the male seemed blacker than the female, but examination of specimens indicate that such is not consistently the case. Foop Fernando Orriz, an Ecuadorian student in our party, called us to see a hawk that had just caught a young marine iguana (Amblyrhynchus cristatus) and carried it to a dead mangrove where it was eating it. The reptile was perhaps 40 centimeters long and presumably from the pre- vious year’s hatch. The hawk consumed everything except the tail and a portion of the entrails. Orriz said that the hawk had not carried off the reptile in a clean swoop but had rather flopped about grappling with it; somewhat below the surface of a large lava fissure. The hawk failed in the first attempt, then tried again and was successful (very likely with another individual iguana for there are hundreds of them on this rocky point). During our stay on Punta Espinosa the egg-laying period of the iguanas commenced. The females trudge about on the sandbars and eventually begin to dig a nesting burrow. On the morning of February 3 as We were watching half a dozen or so thus oceupied, an adult hawk came flying by. Immedia- tely intrigued by this display of exposed iguanas away from the lava to which they can cling, it lit on the sand among them, then ran about four meters and seized one by the back with both feet. The presumably painfully astonished iguana rushed away, dragging or carrying the hawk. The latter, evi- dently accustomed to carrying its prey rather than the reverse, released its grasp and flew to a lava outcrop a few feet away. After surveying the situation, it flew off. For a Source : MNHN. Paris NOTES ON THE GALAPAGOS HAWK 13 photograph of a hawk (probably one of the pair 1 studied) with an iguana as prey see LEVEQUE (1963). Both of these instances suggest that the marine iguanas even when young are at about the upper limit of prey size for the hawk, and are handled somewhat gingerly. That the observed instance of predation was not an isolated one, is shown by the remains of a young marine iguana found in the nest of the hawk and another on lava some distance away. Dr. Herndon DowLin@ of the New York Zoological Society has assumed that this hawk also preys upon the young of the land iguana, Conolophus. There is no reason to doubt this, although, unlike its marine relative, the land iguana bites viciously. Nevertheless, it is a rather clumsy creature and a young one would probably have little chance of sei- zing a hawk. Ross KIESTER, a herpetology student at the Uni- versily of California, told me that he tied the legs of an adult land iguana (on the uplands of Fernandina, their only real stronghold today) to immobilize it. When he returned to pick it up, a hawk was perched on the ground beside it, but had not attacked it as yet. Such behavior, of course, might be expected in almost any predator, quick to observe that a crea- ture is in distress. On Santa Fe Island there is a small surviving population of land iguanas, composed mostly of old adults. Young indi- viduals are very seldom seen. This island is infested with feral goats which have greatly thinned out the vegetation. Hawks are common. DowLinG (1964) has suggested that under such cireumstances the hawks may catch enough of the young iguanas exposed by the grazing of the goats, to lead to the extirpation of the species. This is possible, but it seems likely that the goats might well have the same result eventually, even if hawks were absent, At any rate, all agree that it is the goats, not he hawks, that should be removed. Among the debris about a hawk nest was a skull and other bones of a full grown Galapagos Green Heron, Buto- rides sundevalli. This species is slightly larger than the mainland B. virescens and B. striatus and is another indica- tion of the fact that Buteo galapagoensis, like so many of the birds of these islands, utilizes an extraordinary range of food. Whether the scarlet crabs, Grapsus, which are so superabun- dant along the shore line, are utilized by this hawk is still Source : MNHN. Paris 14 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE not certain. Beneath the feeding perch where the hawk ate the iguana, there were scattered pieces of the shell of a crab or two on the sand. We thought they had probably been eaten by the hawk. On January 30, during a day spent on the high ridges above Tagus Cove, Isabela Island, one or both of a pair of hawks was in view much of the time. One of them was observed to hunt in a method that is very common in Buleo polyosoma and B. albicaudatus. This is to pause or hover at a considerable elevation, then, if no prey is discerned, to glide over to a new station 100 meters or so away and repeat the careful inspection of the ground beneath. This seems very definitely to be a method of systematically examining the ground below for food. The distance which the bird tra- vels before again hovering is more or less the same each time. Thus this method of hunting differs from the hover- hunting of the American Kestrel (Falco spaverius) in which the bird, quite definitely, stops Lo hover only after it spies something interesting on the ground below. In the instance mentioned, though 1 have used the word « hover », a stiff breeze was blowing and the hawk merely hung motionless without flapping its wings. Even so, the wind was strong enough so that the bird’s tail was not infre- quently flipped above its back. The hawk in question was hunting at what seemed to be a rather remarkable height above the semi-arid, sparsely brush-covered slope below, perhaps 50 meters. One could scarcely expect it to see locusts at this eleyation. Lava lizards (Tropidurus) were present though scarce, presumablÿ because of a prevalence of feral domestic cats and may have been the prey sought. On Punta Espinosa, a hawk was seen car- rying in flight one of the endemic snakes Dromicus, which are common in the area. VoicE The usual ery of alarm or excitement is a loud, harsh scream « keer » uttered several times in succession and occa- sionally hurried so that it becomes almost a whinny. The voice is thus not unlike that of Buteo lineatus but is rather different from that of such species as B. jamaicensis and B. swainsoni which utter longer screams singly. Source : MNHN. Paris NOTES ON THE GALAPAGOS HAWK 15 During coition a softer ery, kirp, kirp, kirp, or klilp, klilp, klilp, audible for at least 75 meters, is given continously. I thought only one sex did this, but as noted elsewhere, could not be sure. NESTING HABITS When we arrived on Punta Espinosa, Fernandina Island just at dark, a hawk was perched on a dead mangrove out on the point. It flew away to roost in the gathering darkness. It returned to the same perch early the next morning. This was an immature bird which together with a nesting pair of adults seemed to comprise the population of the species on this part of the island. The immature bird was never seen with the adults except on one occasion. Then, a third bird, presumably this immature, joined the soaring pair of adults for a few minutes before veering off. It is perhaps worthwhile to summarize our day by day observations of the breeding pair, as an example of an inter- rupted breeding cycle. January 26. — As I entered a clump of mangroves 50 meters from camp to store film, I became aware of the pair of buteos cireling just above, screaming. I emerged to watch and saw them cireling about, from time to time drop- ping into the mangroves, once one with legs dangling. Then one landed on an exposed perch, on top of a mangrove. The other followed. The male, noticeably smaller flew off. A few seconds later the male returned and copulation took place immediately. A one-syllabled call, kirp, kirp, kirp, softer than the usual seréam, was uttered continuously, but neither on this or later occasions was I able to tell by which bird. During the morning of this day, Dr. A: H. MizxEr said he had visited the nest, which C. C. CARPENTER of our party had found in use two years before. The birds had protested vigorously. Sr. Ortiz also had visited the nest and again the old birds had reacted strongly. In the aflernoon I looked for the nest and found it to be about a kilometer from camp. It was on the first ridge of lava, where the smoother lava near the shore gives way to very rough, sharp lava that leads away at a slight slope toward the main volcano which com- prises Fernandina. Built of white, barkless, bleached man- 2 Source : MNHN. Paris 16 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE grove sticks from shore drift, the large nest was very cons- picuous. The nest twigs spilled over somewhat onto a lower level of lava. Except for the extremely rough, unstable footing, the nest is easily reached. I found one egg in the nest. It was completely covered by two or three fresh sprigs of mangrove. I had the nest in view from about 1:30 to 2:15 p. m. but saw nothing of the parents. It was very hot, and I wondered if the egg could resist the heat even though covered by the mangrove leaves. However I concluded from the behavior of the birds that the egg had probably been laid very recently. I did not see the birds again all day, but they were observed mating on at least two occasions by others of the party. The nest, when first examined, appeared as though the two or three rather short leafy twigs of mangrove had been placed directly over the egg thus concealing (or shading) it. It did not look as though the green leafy sprays had been merely dropped on the nest, though this seems usually to be the case in those numerous species of hawks that bring such green material to their nests. The extent, if any, to which hawks « deliberately » cover their eggs requires investiga- tion. If the habit exists it has rarely been mentioned by natu- ralists. Yet A.C. BENT (1937, p. 323) wrote as follows of a Bald Eagle’s (Haliaeetus leucocephalus) nest he examined : « No eggs were visible, but I found them deeply buried under fully 2 inches of the soft lining, completely concealed ; the eggs had evidently been covered by the eagle when she left the nest. » I have spoken with individuals who have examined numerous nests of this eagle without noticing such behavior. In a recent note on Parabuteo unicinclus, a close relative of Buteo, it was stated of the three eggs : « These had been covered with strips of inner bark, presumably by the parent bird which was not incubating at the time » (PARMELEE and SrEPHENS, 1964). One of these eggs later hatched, so it is known that the nest had not been deserted. With hawks there is, of course, always the possibility that an incubating bird will inadvertentiy drag some of the nest material over the eggs as it leaves the nest. To revert to the Galapagos Hawk, 1 doubt that the eggs could stand exposure to the sun during the heat of the day for any length of time. Even though incubation in some hawks is said to begin with the laying of the first egg, cove- Source - MNHN. Paris NOTES ON THE GALAPAGOS HAWK 17 ring of the egg or eggs with green-leaved twigs and/or nest lining would have adaptive value, Whether such a habit does in fact exist in any hawk requires confirmation. January 27. — Orriz and I worked our way slowly down to the hawk nest, busied with other observations on the way. Reaching the nest about 9 : 45, there was still the one egg, covered by the leafy mangrove twigs. When I lifted them, the leaves were wet beneath. This had kept the egg cool although by now the sun was becoming warm. Ortiz called my attention to one of the hawks sitting in the top of a dead mangrove by the shore, perhaps 100 meters away (this proved to be a favorite perch). A mockingbird, Nesomimus trifas- ciatus, sat on another dead branch 3 or 4 meters away. They often mob the hawks (if it is possible for one or wo indi- viduals to « mob »). Later, as we walked down the shore, a hawk, perhaps the same one, flew by and screamed once or twice. It occurred to me this morning that the nest may have been deserted, unlikely as this seems. This hawk is, like most other Galapagos creatures, very lame. But that does not mean thal, if the nest is visited on several occasions on the very morning when the first egg was laid, the parents might not desert. SwarrH (1935) pointed out that the Gala- pagos dove, Nesopelia galapagoensis, has a strong distraction display even though, he thought, it has been superfluous ever since the dove’s ancestors reached the islands, some tens or hundreds of thousands of years ago. Thus behavioral traits, strongly embedded in the genotype, are perhaps sometim relained almost indefinitely, just as are physical traits (May, 1963, p. 288). A. H. Mixer however doubted that the egg was fresh and pointed out that it was nest stained. However, the midday sun literally cooked the juice from the fresh mangrove leaves above it, and this may have stained the egg quickly. 1 did not see the hawks again on the 27th, but Professor CARPENTER saw one of them swoop down and carry of a young marine iguana. January 28. — Visit the nest at 8 a. m. Already the sun is getting hot. The mangrove leaves in the nest are now quite wilted. Am now convinced the nest is deserted. One of the hawks, however, perched for a few minutes on the high Source : MNHN. Paris 18 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE branch near the shore, then flew silently down into the man- groves. January 29. — As I walked down the foreshore towards the nest, one of the hawks flushed from the ground. It flew to a perch about two meters high. A mockingbird flew to a nearby perch and scolded once. I reached the nest a little later at 10 a. m. Just as I was examining it, a couple of weak screams called my attention to one of the hawks on the high mangrove perch. Apparently the birds, or one of them, retain a mild interest in the nest. The now dry mangrove leaves and twigs which once covered the egg had blown away. 12 : 30 p. m., Male hawk diving and screaming, as female perches in a tree. 1 p. m, Pair high in air, facing motionless into breeze ; one or both screaming. Soon one, presumably male, cireled higher, then descended towards soaring female, with legs dangling. Both soon glided of and eventually disappeared behind a clump of mangroves, screaming loudly. It would seem the pair, after abandoning nest, is coming into the active mating phase of the cycle again. January 30. — (Absent from area). January 31. — Visited nest. Hawks not present. Egg still in nest, One hawk seen later in morning catching and eating young marine iguana. February 1 and 2. — (Absent from area). February 3. — 8 : 30 a. m., The two hawks fly to adjacent perches in a dead mangrove. Both preen and clean their beaks. 8 : 38, Female flies to another snag 50 meters off ; soon drops to a lower limb, legs dangling and perhaps soli- citing male who continues to perch stolidly. 8 : 40, Female leaves perch ; when she has covered 5 meters or so, the male leaves his tree and follows, screaming loudly. They are quickly lost from sight, but a little later I observe them screaming and plunging in the distance. 9:55, Both hawks back on original perches. Male soon departs. Female remains on a half concealed perch, occasio- nally changing directions. Perhaps she is looking for prey, but the low mangroves below are so dense it seems a poor Source : MNHN. Paris NOTES ON THE GALAPAGOS HAWK 19 hunting place. 10:40, Male appears silently and perches about 3 meters from female. After a few seconds, he hops to perch half that distance away, then to side of female and coi- tion follows at once. The male soon flies off, followed by the female. They soar and flap about, with some screaming. The female returns to perch in the same tree, where she screams occasionally. A little later I see three birds circling high, at least one of them screaming ; but one of them (probably the immature that is in the vicinity) soon scales off by itself. (Later this morning, one of the adults, probably the female, had the encounter with nesting iguanas described earlier.) February 4. — 1 p. m. One hawk perched on highest branch of tall, dead mangrove. Flies off behind some trees, but a minute or so later, two hawks come flying back to same perch and copulate immediately, one or both uttering a high pitched, repeated Xlilp, audible when I am, perhaps 100 meters away. Both preen, the female more vigorously. 1:10, Again copulate, then resume preening. Five minutes later the female hops to a perch close to the male. A few minutes later she fans her tail feathers, stretches one wing and turns towards her mate. 1: 30, The female flies off ; and when I look a minute or two later, the male has gone also. ++ The remarkable fact to me was that the pair had resumed active sexual behavior without being seen near the nest, which still held the deserted egg. Possibly, however, the female would simply have laid in the nest, without further preliminaries, except, possibly, to push out the old egg. It was considered unlikely that the pair had another nest, as our party scoured the entire area rather thoroughly. Lack (1950, p. 273) considered the Galapagos buteo a breeder during the « cool, dry, season » and found records of eggs from April through August. LEVEQUE (1964) found that with increased data the nesting season of many Gala- pagos birds, including the hawk, is more prolonged than was Source : MNHN. Paris 20 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE previously believed. He knew of egg records for Buteo in April, June, August, and December. We have added January and probably February (pair mating, Santa Fe Island). During the same period a pair observed on Española, showed no evidence of nesting beyond the fact that the birds tended to stay together. The ecology, including climate, varies tremendously on the islands. Even on a single island, one goes from semi-arid lava desert to wet highland forest. In some years there is more rain than in others. The hawk is found in a variety of habitats. Further studies may show, however, that there are local regularities in the season. Especially on low, relatively unvaried islands such as Española, the hawks may have a definite season, perhaps correlated with cycles in some staple food. Or it may be that there is no definite season of repro- duction and that nesting is on some non-annual cycle for each pair, or is irregular and triggered by the food supply. RESUME Monographie de la Buse des Galapagos (Buteo galapagoen- sis). Position systématique et comportement. Cette seconde par- tie fait état des résultats de la mission patronnée par la Natio- nal Science Fondation et l'Université de Californie de Ber- keley. L'auteur consigne ses observations personnelles, faites en janvier, février 1964, sur cette espèce jusqu'ici mal connue : nourriture, voix, reproduction. Sur ce dernier point, nous trouverons une série de notes particulièrement intéressante, prises quotidiennement au cours de la reproduction d’un couple sur l'Ile de Fernandina : copulations, nids, œufs. Malheureusement cette observation n’a pu être menée à bien jusqu’au bout. LITERATURE CITED Bent, A. C. 1937. — Life Histories of North American Birds of Prey. U. S. National Museum, Bulletin 167, 409 pp. Bnosser, A. 1963. — Le comportement de la Buse des Galapagos Buteo galapagoensis. Alauda, 31, pp. 5-21. Source : MNHN. Paris NOTES ON THE GALAPAGOS HAWK 21 Dowiinc, H, G. 1964. — Goats and hawks : a new theory of predation on the Land Iguana, Animal Kingdom, April, pp. 51-56. JoHxson, N. K, and H. J. Pa 1963. — The systematie position of certain hawks in the genus Buteo. Auk, vol. 80, pp. 417-446. Lack, D. 1950, — Breeding seasons in the Galapagos. Jbis, vol. 92, pp. 268-278. Lévêque, R. 1963. — Le statut actuel des vertébrés rares et menacés de Varchipel des Galapagos. La Terre et la Vie, 4, pp. 397-430. — 1964. — Notes sur la reproduction des oiseaux aux îles Galapa- gos. Alauda, 32, pp. 5-44; 81-96. May, E. 1963. — Animal species and evolution. Cambridge, Massachu- setts, Belknap Press, Harvard University Press. Panmeee, D. F., and H. A. Srepmexs. 1964. — Status of the Harris Hawk in Kansas. Condor, vol. 66, pp. 443-445. Ripeway, R. 1873. — Catalogue Ornithological Collection Boston Society. I. Falconidae. Proc. Boston Soc. Nat. Hist., vol. 16, pp. 41-106. — 1875. — On the buteonine subgenus Crazirex, Gould. Proc. Acad. Nat, Sci. of Philadelphia, pp. 89-119. Swanrs H. S. 1935. — Injury-feigning in nesting birds. Auk, vol, 52, pp. 352-354. Source : MNHN. Paris SOME REFLECTIONS ON TWO RECENT VISITS TO THE AZORES ARCHIPELAGO by David BANNERMANN, SC.D., LL.D. Honorary Associate, British Natural Historg Museum At 7 p. m. on the 29th May 1963, the Portuguese mailboat CARVALHO ARAUJO, due to have sailed four hours earlier, edged her way out of Funchal harbour and set her course for the Azores, a group of islands to which I had always hoped fate would some day bring me, though my hopes were not to be realised until the eventide of life. It was already begin- ning to get dusk as we passed the great cliff of Cabo Giräo rising close on 2,000 feet sheer from the sea, from the sum- mit of which we had once watched several huge sting-rays (Trygon pastinaca) in the clear waters at its foot. As the din- ner bell sounded we were approaching Ribeira Brava and when later we came up on deck the daylight had gone, the twinkling lights from the villages on the mountain side were all that remained, and the flashing beams of the lighthouse on Ponta do Pargo was the last we saw of Madeira as the CaRvALHO passed from the shelter of the land to the open sea. The weather was all it should be and the voyage to Santa Maria, the first island to be visited, was over almost before we had realised our trip had begun. It was our intention on this occasion to visit as many of the nine islands which com- pose the Archipelago as cireumstances would allow, for lan- ding and re-embarking even in a normal sea is not to be lightly undertaken where there is no harbour, and when the rise and fall of the waves is always considerable. Only at Ponta Delgada in Säo Miguel and at Horta in Faial are there really first rate harbours, in all the other islands, Santa Maria, Terceira, Graciosa, Säo Jorge, Pico, Corvo and Flores, the vessels loading or unloading stand out to sea and passengers — as we were warned — go ashore at most of the little ports L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris RECENT VISITS TO THE AZORES ARCHIPELAGO 23 < at their own risk ». We were to consider ourselves very lucky that only on Graciosa and Säo Jorge was it impossible to go ashore. In the course of her leisurely journey around the islands the CarvaLHo AnauO visited each island twice, both on the outward journey to Flores and on the return. Sometimes the visit was for the whole day or two days according to cargo, and sometimes less, but it gave us the opportunity to obtain a much better idea of the bird popula- tion than such brief visits would suggest. À fascinating account of these islands and their bird life could be written by a visiting naturalist with a gifted pen for description, but in this short article in which I was restricted to « five pages » of L’Oiseau et la R.F. 0. only the high lights of our voyage can be mentioned, and only general impressions given. We have returned again in the autumn of 1964 to Säo Miguel, where we have stayed another five weeks, having set our- selves the problem of discovering if the priôlo — the endemic bullfinch of the island — still exists anywhere in the remotest forests, but of that more will be said later. 1 was already familiar with Azorean literature and many years ago in the British Museum had made a study of the spe- cimens collected in the Azores by F. Du Cane Godman, and later by W. R. Ogilvie-Grant, whose published reports on their collections — remarkably complete ones so far as the resident birds were concerned — were the foundation upon which others following afterwards have built. On this first voyage, accompanied by my wife, we were to form some very definite impressions of the status of the Azorean birds and of the work which remains to be done, for field work, as apart from collecting for which there is no more need, has been sadly neglected. The first thing which strikes the visi- ting naturalist is the paucity of resident land birds, and the number of species which one might expect to find, which have never occurred. A few species are dominant and wide- spread and these can be counted on the fingers of one’s hands ! Chaffinches, only one race occurs (Fringilla coelebs more- let), abound everywhere, and of the other Fringillidae ; Canaries (Serinus canarius) are equally numerous. Grey Wag- tails (Motacilla cinerea patriciae were seen on every island from the coast to the higher elévations, Redbreasts (Eritha- cus rubecula), Blackbirds (Turdus merula azorensis) and Blackcaps (Sylvia atricapilla atlantis) were constantly in evi- Source : MNHN. Paris 24 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE dence and wherever the heaths or pines were prominent we were sure to see Goldcrests (Regulus regulus), the only bird in the Archipelago be it noted of which more than one subspe- cies has been described. There are in fact three. All other land birds, except those enumerated above, are relatively uncommon, except perhaps the insular Quail (conturbans) which, owing to the number of eggs produced in a season, and difficulty of flushing, is able to hold its own, while some other species are quite definitively dwindling. One bird of which we saw a number on the western and central islands, but which appears to be absent from Corvo and Flores, is the Buzzard, a small dark endemic race which was named in honour of the late Lord Rothschild (Buteo buteo rothschildi). It is said to have been once very numerous and is believed to have been the bird which the first discoverers of the islands mistook for the Goshawk, the « Acor » of Portugal, after which the islands are named. As no Goshawk (or Kite) has ever been reported from the islands the inference is reaso- nable. Starlings (Sturnus vulgaris granti) which we met with in varying numbers on Santa Maria, S. Miguel, Terceira, Pico, Faial, and Corvo, though not seen by us on Flores, were relatively common. The sub-specific distinction of this Azores starling has recently been questioned by an American. During our travels in the various islands we formed the opinion that not more than ten land birds could be reckoned as really common! The numbers of these was sometimes striking, as in the case of the Chaffinch, Wagtail and Canary, but there are other birds present, though much less in evi- dence : Goldfinches (introduced) and Greenfinches (also reported introduced) and Spanish red-legged partridges which must at one time have been imported from somewhere north of the Douro judging by the brillance of their plu- mage and russet-hued mantles, but these birds are not encountered by the casual visitor, and the finches are both definitely rare and local at the present day. The Azores can claim only one resident Owl and that the long-eared. The few specimens collected show no distinction from mainland Asio otus otus which is rather surprising, as they must have been resident in these Oceanic islands for a great many years and are unlikely to have reinforcements arriving from the Continent. Of the endemic races which must be conside- Source : MNHN. Paris RECENT VISITS TO THE AZORES ARCHIPELAGO 25 red to be nearing extinction the Azorean wood-pigeon (Golumba palumbus azorica) is unfortunately one, while the Azorean bullfinch (Pyrrhula murina of Godman) is close on the verge. This bird and its status in Säo Miguel, if it still existed, was the real object of our return to the Azores in the autumn of 1964, and this seems to be the proper place in this brief article to recount the result of our enquiries. Armed as we were with a coloured plate of the Bullfinch by Mr. David Reid Henry, we were in a better position to tackle local opinion and interview those reputed to have known the bird in olden days than we were during our first visit in June of the previous year, when we searched the slopes of the mountains above the lake of Furnas and the orchards by the little stone-built chapel on the shore whence had come the last examples ever to have been obtained in 1927. The Priôlo — by which name it is generally known — has always been restricted to the eastern provinces of S. Miguel and occurs nowhere else in the world. In the year 1865 F. Du Cane Godman collected what specimens he needed in a poplar tree in the space of a few minutes, the report of a gun did not alarm it, so tame was it reported to be — or so unused to firearms. Its refusal to spread beyond its immediate habitat, the orchards and forests east of Fur- nas, points to its sedentary habits and its attachment to a particular terrain. Its fondness for peach and other fruit blossom is the real reason why it has been reduced to a few pairs. That it still exists in the more inaccessible forest patches my wife and I, after endless enquiries on the spot, and meetings with those who have known it in the past, feel reasonably sure. It was, however, mostly the older men who recognised the coloured pictures of the Priôlo which we car- ried with us, those of the younger generation had obviously no personal knowledge, but that it had been actually seen within the last two or three years by responsible persons of middle age was our final opinion. Search for it as we did in ravines (ribeiras), mountain glades, Cryptomeria forest, orchards, heather-clad slopes, and on private estates where it had been reported as occurring within recent years, we never saw anything which we could definitely assign to the Priôlo. More than once a bird would cross our path to disap- pear all too suddenly into thick growth which had the flight and appearance of a bullfinch, but in no instance were our Source : MNHN. Paris 26 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE hopes fulfilled, and we left the area where it is reported to oceur with the conviction that the only chance of seeing it would be when the orchards are gay with blossom and the birds have vacated their winter retreats, wherever they may be, to feed on the fruit-buds. The total extinction of the Azores bullfinch — when it takes place — must not be attributed to collectors of skins for preservation in Museums, but unquestionably to the islan- ders themselves who took every opportunity to slaughter the birds on account of the damage done to the fruit trees, ridi- culously unafraid of man as it was reported to have been. There are grounds for believing that a reward was offered for every bullfinch beak produced to the local agricultural authorities, and with every man and boy (the latter armed with catapults) on the watch for it, its days of plenty were very soon numbered). As the only endemic species in the whole of the Archipelago, its extirpation will be a tragedy which it seems impossible to prevent. Once a species is redu- ced to such low proportions it is almost impossible for it to recover. Säo Miguel where the Bullfinch has its home is the island in which most bird research has been carried out and to which visiting ornithologists of other nations have been attracted, but interesting though this large island undoub- tedly is, with its crater lakes and varied scenery, the other islands of the group all have their particular charm. In Ponta Delgada, the capital of Säo Miguel, the very excellent Museum «€ Carlos Machado » is a natural rendez-vous for foreign visitors, and is under the capable honorary direc- torship of Senhor José Maria Alvares Cabral. He has already done wonders in re-organisation and the determination of specimens under his charge and by sceing that all newly acquired material is properly recorded and labelled. We were greatly impressed by his ability. The Museum has suffered from having far too many extra-limital exotic species on exhibition, with the result that the cases are all overcrowded. This is a legacy which Senhor Alvares Cabral is doing his best to remedy since he has been in charge. From Säo Miguel the CarvaLHO ARAUJO sel course for Terceira, the first of the five in the central group of islands to be visited. Angra Do Heroismo, of which we anchored, was at one time, prior to 1832, the capital of the Archipelago Source : MNHN. Paris RECENT VISITS TO THE AZORES ARCHIPELAGO 27 and is nearly 100 miles distant from Ponta Delgada, which succeeded Angra as the capital town and principal port. It was here on the 2nd of June 1963 that we met, and spent some time with, a naturalist of wide repute, Colonel J. Agostinho, whose knowledge of the zoology and geology of the islands is second to none in the Archipelago. To his credit was the first discovery of Bulwer’s petrel, Bulweria bulwerü, breeding in the Archipelago, as recorded in < Alauda » 1937, and of the visit to several of the islands in the winter of 1963 of a party of Mute Swans Cygnus olor, two of which fell to gunners in Terceira and were duly pre- served. The first occasion on which any Swan had ever been reported from the Azores Archipelago, Colonel Agostinho reported them from the first as mute swans, but a female specimen, now in the Museum at Ponta Delgada, gave some reason to doubt the identity, and not until photographs of the specimen had been sent to the Smithsonian Institution in Washington, where they were expertly examined by Doctors Alexander Wetmore and John Aldrich, was the identification C. olor confirmed. Female swans are extremely difficult to identify and the possibilily of an American species being involved — as first suggested by Senhor Alvares Cabral — could not be excluded. Terceira in the not so distant past had the ornithological advantage over the other islands in the central group of pos- sessing a good sized marsh in which the Moorhen and other marsh-loving birds were wont to congregate and breed. The moorhen of Terceira has been described as an insular race but its haunts in the island have now been drained and we learned from Colonel Agostinho that it is now seldom scen, though not yet extinct, for a young bird has recently been brought to him. The Coot is another bird whose numbers are dwindling as a resident, but which can look for reinforce- ments from overseas in the winter months, and it was to this marshland that Lille Bitterns from Europe occasionally made their appearance. There had been a small incursion of this species Lo the Azores a month before our visit and one or two had even reached Madeira where we chanced to exa- mine specimens which had been brought to the Museum. It Was during our first visit to Angra that Colonel Agostinho suggested the possibility from its behaviour — as witnessed by himself in Terceira — and from information received from Source - MNHN. Paris 28 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE a correspondent in Graciosa in whom he placed confidence, that the Turnstone Arenaria intérpres occasionallÿ bred in the Archipelago, for news had reached him of a parent suppo- sedly of this species having been seen between Vitoria and the lighthouse on Ponta da Barca « leading its young on the shore to a safe retreat ». As in the course of my long expe- rience in the British Museum and elsewhere I had received similar assurances from other areas in which northern bree- ding species were said to have been discovered breeding far beyond their known range, but which, on further examina- tion, were perforce refuted, I dismissed these breeding records as too unlikely to be possible, feeling some error of identifi- cation must have occurred. Great was my astonishment on returning lo the Azores in 1964 to be met on the ship by our good friend Sr. Cabral, the honorary director of the Museum Carlos Machado, with a photograph of a chick of a shore bird taken by a German investigator in the month of June 1964, purporting to be that of Arenaria interpres ! 1 confess that I was almost convinced but not quite. There was something wrong, the tarsus too long, the eye too large. The markings did not quite agree with those of Turnstone chicks in the Royal Scottish Museum and British Museum, photographs of which I had taken the precaution to bring out with me to the Azores. Scottish caution prompted me to send that pho- tograph to a friend in Britain, who submitted it to the expert opinion of ornithologists on the staff of our two leading Museums. With skins of the chicks of wading birds before them for comparison, the verdict was reached unanimously that the chick captured (but not preserved) on Terceira on the 29th July 1964 at Praia was that of the Kentish plover, Charadrius aleæandrinus. That opinion has now been endor- sed by two well-known naturalists who are familiar with the Turnstone chick in Norway. Thus was exploded yet another myth — just in time! Photographs of birds, unless very exceptional, are a dangerous means by which to make certain identification. It is a lesson which should be taken to heart in this age, when the collecting and preserving of specimens in the aid of scientific research is looked upon as reprenhen- sible, Identification by means of photography alone is bound to lead to errors. Our next port of call was Graciosa and as we neared the island we were delighted to see in our wake the first Stormy Source : MNHN. Paris RECENT VISITS TO THE AZORES ARCHIPELAGO 29 petrels — almost certainly Oceanodroma castro, which we had met with on this voyage. We had already made history by identifying an Arctic Skua, Stercorarius parasiticus, some five miles out from Angra, believed to be recorded from the Azores for the first time, and had passed close to some high larva cliffs on the ledges of which Common Terns, Sterna hirundo, were unquestionably breeding, the first time we have ever seen {his tern nesting in such a situation, although we have heard of it doing so in the Desertas off Madeira. As the CARVALHO passed, the terns, for some reason, panicked and, accompanied by hundreds of Rock-pigeons, Columba livia atlantis, poured from the ledges, transforming the dark eliff face into a whirling mass of while and blue — a very lovely sight. It was not until 3 p. m. that our ship dropped anchor off the little port of Praia, and as it was pouring with rain and we lay far from the landing place, we elected to remain on board. Our time was not wasted, however. Lying off shore was a low-lying island from which a continuous stream of terns were making their way to fish in a shallow bay, upon the shores of which tiny waves were breaking. As the birds passed to and fro we observed that on their return journey their bills were filled with little silveryÿ fish, a sure sign that they were feeding young. Closer inspection revealed that all but a very few were Roseale Terns, Sterna dougallii, making for the low sandy spits which jutted out to sea at either end of the island. There must have been hundreds of them — the largest roseate colony which either my wife or I had ever seen ; bird after bird passed close Lo our ship, allowing an unrivalled view of their bills, the greater part of which were black, and of the exceptionally long streamers in the tail ; we were to meet with other roseate colonies again in Faial, where they were in the minority, but on this islet off Gra- ciosa the roseates held complete sway. When we realised how seldom they had figured in the Azorean literature — Godman had recognised a few amongst a common tern colony off Faial in 1865, and José Correia had collected a spécimen in 1927 — we were all the more astonished by the numbers we encoun- tered. If, as seems improbable, they can remain unmolested by local egg-thieves, it will be one of the largest breeding places of this tern in the Western Atlantic. Roseate terns are Proverbially fickle however, here today and gone tomorrow, Source - MNHN. Paris 30 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE for very little disturbance will cause them to leave their bree- ding place, as we know to our cost in the British Isles. From Graciosa our voyage took us to Säo Jorge where we anchored after dark under the shadow of tremendous cliffs to land passengers and leave again before dawn. Daylight found us lying off à tiny port named Prainha on the north coast of Pico, where we were fortunate to find a car in which we traversed nearly the whole of the island before laking a fast launch at Madalena in which to cross the dividing strait to Horta, the capital of Faial. Pico is a delightful island well covered with trees, the rich larva soil planted with crops and vines. The roads were bordered with white and yellow daisies (a refreshing change from the ubiquitous hydrangea), the brilliant pink of the ice- plant (Mesembryanthemum) and many coloured geraniums. The « Pico » itself rises steeply to 2540 metres (7613 feet) and is often covered in cloud. Great larva flows run from its cone to the sea and forest covers the lower slopes of the mountain, the former haunt of the Azorian wood-pigeon (Columba palumbus azorica), which is now a scarce bird eve- rywhere. All the commoner birds were seen on Pico but the pigeon — if it still exists — remained hidden in the woods where it was so numerous in days gone by. Of our experiences in Faial we have no space to write __ the island interested us much less than the next two to be visited : Corvo and Flores, lying farthest out in the Atlantic, isolated by more than 100 miles of sea from Faial and Gra- ciosa, and 680 miles east of the Grand Newfoundland Banks and the nearest American land — these outposts in the Ocean are to my mind the most romantic of all the islands we had visited. In Corvo, upon whose treacherous rock-girt shore we landed on the 5th June 1963, one might have stepped back 100 years, so primitive did life appear to be. In Corvo and Flores, the latter rightly famed for the beauty of its scenery and the luxuriant vegetation it supports, much of it, alas, introduced from warmer climes, we have the two islands first to be sighted by American vagrant birds, of which there is now such an imposing list. Corvo is said to be the only island in the Archipelago upon which the Manx Shearwater (Puffinus puffinus) breeds, but that can be disputed. It is the smallest island of the group, 4 miles ‘/, by 3 miles, with its population of under 1000 gathered into the only township, Source : MNHN. Paris RECENT VISITS TO THE AZORES ARCHIPELAGO 31 Rosario, with its highly precarious landing place amongst jag- ged and terrifying rocks sticking up at every angle. Our lan- ding there was something Lo be remembered and only possible on what the islanders — and probably no one else —— term a calm day. Flores « where Sir Richard Grenville lay » is only a short, distance from Corvo and as we neared the island rafts of Cory's Shearwater (Puffinus diomedea borealis) — the < Cagarra » of the Portuguese — were seen floating on the sea, hundreds of birds in every « raft ». A line of huge lighters filled with large black and white cattle, drawn by a small launch, came out to meet the CARVALHO, for there is a regular trade of these draught animals between Flores and Lisbon. The work of hauling them aboard from the tossing lighters was a sight we shall not lightly forget. The boatmen of Corvo and Flores are renowned for their prowess ; they must be second to none in the world for the skill with which they manage their boats in turbulent seas. Flores impressed us tremendously. A drive half way round the island by a road cut out of the cliffs before turning inland and climbing to the central moorlands enabled us to obtain a good idea of its varied scenery and bird life. The high ground on this island reminded us of a Scottish moorland and we could well imagine that Teal, Mallard, and Snipe would bring up a family in such favourable surroundings if permitted to do so. From one point it is possible to leave the road and look down upon à beautiful crater lake upon which two terns were fishing. Fine as is the inland scenery of Flores, that of the coast is grander still The coast line between Santa Cruz and Lajes das Flores is truly magnifi- cent. Here, if anywhere, the dark Rock-pigeons which I named atlantis are able to breed true. Land birds on these islands so far distant from their nearest neighbours —— some 150 miles — have not, as one might expect, become differen- tiated from those in the central group of islands, but in Flores at least they were well represented by the usual ubi- quitous species. In this short essay 1 have attempted to give my impres- sions of the Azores as seen by a visiting naluralist with no special qualifications for the job other than my interest in Oceanic islands. The Azores are typical examples of such, always poor in the number of breeding birds and in this ins- 3 Source : MNHN. Paris 32 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE tance not in the course of any regular stream of migrants, as is for instance Cyprus or the Canary Islands. Even so, the Azores are remarkable for the number of migratory birds, wind-blown or sea-borne vagrants, which have reached their shores. One hundred and twenty-five species have already been recorded in this category. While the great majority are of European origin, no less than twenty-one of the vagrants bail from North America, representatives of {he Families Fregatidae, Podicipitidae, Ardeidae, Alcedinidae, Cuculidae, Rallidae, Anatidae, Charadridae, and Turdidae, the last named contributing the only American Passerine species to have crossed the 2000 miles to reach Säo Miguel. The Ame- rican influence has never been sufficiently stressed but it was mentioned by Staffan Ulfstrand in his interesting obser- vations « On the Vertebrate Fauna of the Azores » (Bol. Do Museu Municipal Do Funchal, XIV, 1961, pp. 75-86) quoting only 10 species of American origin known to Mayaud and De Chavigny in 1932. Since then the list has been more than doubled. Ulfstrand points in his short review to the north- central European character of the Azores avifauna rather than to a south European, caused he believes by the stron- ger migratory tendencies of the northwest European popula- tions. One species of Arctic affinities must receive special men- tion : the Snow Bunting (Plectrophenax nivalis). It may now be reckoned a regular passage migrant to the Archipelago. It has visited the islands in small numbers since the days when F. Du Cane Godman reported it from Corvo in 1864-65, to which island Senhor Fernando Rocha, Chef du Service Météorologique, now considers it an annual visitor in winter. We have seen it ourselves at sea off Flores when travelling on a ship in April bound from Chile to England via Panama and Curaçao. Another link with Greenland is furnished by the arrival at intervals of the Greenland Wheatear, Oe. oe. leucorrhoa ; of the seven specimens in the Carlos Machado Museum at Ponta Delgada of Oenanthe oenanthe, every one proved by its measurements to belong to the Greenland race. No example of the typical subspecies is in the collection. In Madeira, on the other hand, the only Wheatear collected on the main island is Oe. oe. oenanthe. On Porto Santo specimens of Oe. oe. leucorrhoa have been obtained. I have examined and Source : MNHN. Paris RECENT VISITS TO THE AZORES ARCHIPELAGO 33 measured all these birds. We are still at a loss to account for Godman’s extraordinary statement (Natural History of the Azores, p. 25) that he found « four or five pairs (of Oe. oenanthe) in the old crater on Corvo which had bred there, as I saw young birds that could scarcely fly »… That visit of two days duration took place about the second week of May 1865, he did not collect a specimen on Corvo. AS earlier remarked in this essay, the number of land and fresh water birds known to breed or to have bred in the Archipelago is strikingly small, for they have nine islands from which to choose, covering a huge area of sea. True land birds do not exceed twenty species in all which are known to breed regularly. This number can be raised to thirty-five if we include the summer visitors, the sea birds and the petrel family (Procellariidae) which come to these islands to breed, the ducks (only two), and marsh-birds which breed only exceptionally under favourable conditions and are invariably harried by man. Of the resident species at least three have been introdu- ced — the Partridge, the Goldfinch, and the Greenfinch, and now of all birds the Common sparrow, Passer domesticus ! Strange indeed is the absence of all Birds of Prey (Falconi- dae) except for the Buzzard, all Warblers (Sylviidae) except for the Blackcap, all Swallows (Hirundinidae) and all Swifts (Apodidae). There are no Woodpeckers and only one resident Owl ! We believe that we saw an un-recorded bat, considera- bly larger than Nyctalus azoreum, the only known species. I have already over-run the length allowed me for this essay, for which I must beg forgiveness. Likewise, 1 must crave the indulgence of those who may read these notes, and especially my old and valued friend Jacques Berlioz, in whose honour his friends are contributing to this number of L'Ois. et la R.F.0. for I am well aware these scattered observations are not worthy of so important an occasion, They have, howe- ver, been written under some difficulty in Madeira immedia- tely on our return from the Azores with impressions still fresh upon me. May I end by expressing the hope that our honoured friend and colleague, Professor Berlioz, may have many happy years in which to enjoy his retirement and pursue his travels. He carries with him, wherever his steps may take him, the warmest regards and sincere good wishes of his Source : MNHN. Paris 34 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE many friends, not least in admiration, the writer of these notes. RESUME Relation ornithologique d’un voyage effectué en mai- juin 1963 aux Açores. Voyage au cours duquel toutes les îles de l’Archipel ont été visitées, à l'exception de Graciosa et de Sao Jorge restées inaccessibles, l’état de la mer et l'absence de port interdisant tout débarquement. Malgré d’actives recherches, l’auteur est obligé d’avouer qu’il lui fut impos- sible d’observer les deux espèces qu’il avait placé en tête de liste : le Ramier des Açores, Columba palumbus azorica, et le Bouvreuil des Açores, Pyrrhula murina. La première semble de plus en plus rare, quant à la seconde, il faut craindre, d’après les informations recueillies, que lon doive désormais la classer parmi les espèces éteintes ou bien près de l'être. L'auteur redresse une erreur d'identification qui avait laissé supposer dans un récent passé que le Tournepierre, Arenaria interpres, aurait niché aux Açores. L’invraisemblance de l'information l’incitait à vérifier les faits de plus près. Pru- dence justifiée, car il s'avère que l'oiseau signalé comme entouré de ses jeunes n’était. qu’un Pluvier de Kent Cha- radrius alexandrinus. Par ailleurs le D' BanNERMAN signale une importante colonie de Sternes de Dougall, Sterna dou- galli, à Graciosa, et précise que 4 espèces ont été introduites : la Perdrix rouge, le Chardonneret, le Verdier et malheureuse- ment le Moineau domestique. Pour conclure, l’auteur souligne la rareté des Rapaces (à l'exception de la Buse), des Sylviidés (sauf la Fauvette à tête noire, Sylvia atricapilla) et, chose curieuse, des Hirondelles et des Marlinets. Source : MNHN. Paris AN ANALYSIS OF BIRD CASUALTIES ON THE ROADS IN THE SOUTH WESTERN CAPE PROVINCE, SOUTH AFRICA by G. J. BROEKHUYSEN Department of Zoology, University of Cape Town INTRODUCTION Information on bird road casualties in Britain has been published by Finnis (1956, 1960), and Hopson (1959, 1960, 1962) ; in France by Finnis (1959, 1960) ; in Holland by Môrzer BRuINS (1959) ; in Germany by BEckmanx (1961), Hezn (1961) and Martens (1962) ; in America by Wire (926, 1927), LinspaLE (1929), RoBerrsoN (1930), BARNES (1936) and ScHorGer (1954). No information on bird casualties on South African roads has been published. Since the last war, roads in South Africa have undergone a remarkable development and many of the main roads are now tarred. This has resulted in increase of traffic and also in speed. From April 1960 until the end of March 1962 I gathered information on bird casualties on the roads in the South Wes- tern Cape Province roughly within a radius of 250 miles from Cape Town. During this time any mileage I covered in my car during the daytime was noted and any bird casualties encountered were recorded. In addition to my own observa- tions others were received from a number of members of the Cape Bird Club (S. W. Cape Branch of the S. A. Orni- thological Society). I am very much endebted to these contri- butors of which the following should especially be men- tioned : Dr. J. M. Winterbottom, Messrs. R. K. Schmidt, B. M. Mackenzie, J. Martin, J. G. R. MacLeod, J. H. Hofmeyr, G. D. Underhill, Mesdames M. K. Rowan and I. Taylor and Misses E. B. Robinson, J. Robinson and C. St. C. Robinson. The combined observations over both years cover a total of 41500 miles. L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source - MNHN. Paris 36 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE RESULTS 1) AI the recorded casualties have been tabulated in Table 1, where they have been broken down into species and diffe- rent months of the year. In addition the number of miles for each month has been indicated. The table shows that during the two years concerned as many as 72 species of birds were recorded as road casualties. À number could not be identified (a) because they had been dead for a considerable time and were flattened beyond reco- gnition by traffic, or (b) because due to cireumstances they could not be closely examined nor collected for future exa- mination. During the total period of observation of two years and a recorded mileage of just over 41500 miles, 584 bird road casualties were recorded. umber of birds and number of species killed 2) Comparison between the breeding-season and the non-breeding-season The breeding-season in the South Western Cape Province is from August to January and the non-breeding-season from February to July. There are of course certain exceptions such as the Sugarbird (Promerops cafer (L.)) and the Orange- breasted Sunbird (Anthobaphes violacea (L.)) which are both chiefly winter breeders. The Cape Thrush (Turdus oliva- ceus L.) has a secondary breeding peak in the autumn and quite a number of species of duck nest in July. These few will, however, not effect the general picture. Differences in road casualty rate between breeding- and non-breeding-season can be expected to occur. In Table 2 the the number of casualties and mileage covered has been sepa- rately tabulated for the breeding- and the non-breeding- season. In both years the number of casualties in the non- breeding season is considerably lower than in the breeding- season. The number of miles covered differs for each of the two seasons in each of the different years. The difference for 1960-1961 is 5913 miles and for 1961-1962 it is 676 miles. Source : MNHN. Paris BIRD CASUALTIES IN THE CAPE PROVINCE 37 If the results for the two seasons are being compared the mileage should strictly speaking be about the same. If one assumes that the number of casualties is proportional to the distance covered one can calculate what the casualty figures theoritically should have been if the mileage covered in both seasons would have been the same. TABLE 2 Number of casualties and mileage during the breeding-season (August-January) and non-breeding-season (February-July) Number Number Miles per Season of of casualty casualties miles 1960-1961 April-July 1960; February, March 1961 74 1620 103 August-December 1960; January 1961 243 13533 55.7 1961-1962 April-July 1961; February, March 1962 76 9839 129.5 August-December 1961; January 1962 192 10515 548 1f this adjustment is made the following figures are obtai- ned : 1960-1961 : Non-breeding-season 133, Breeding-season 243 casualties. 1961-1962 : Non-breeding-season 84, Breeding- season 192 casualties. It seems, therefore, that in each of the two years bird casualty figures during the breeding-season were about twice as high as during the non-breeding-season. 3) Species most frequently killed From Table 1 the total number of casualties of any one of the 72 identified species for each of the two years and for the total two year period can be calculated. The eight most frequently killed species are tabulated in Table 3 and pic- tured in Plates I and Il. From this table it can be seen that the order based on frequency differed somewhat for each of the two years. During the period 1960-1961 the Bokmakierie headed the list of casualties closely followed by the Cape Sparrow and then the sequence was Laughing Dove, Fiscal Shrike, Turtle Dove, Cape Canary, Cape Weaver and Spotted Eagle Owl. For the period 1961-1962 the Cape Sparrow headed the list, closely Source : MNHN. Paris 38 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE followed by the Cape Weaver and the others followed in the sequence Fiscal Shrike and Cape Canary, Cape Turtle Dove, Bokmakierie, Laughing Dove, Spotted Eagle Owl. TABLE 3 Species which were most frequently killed Species Number of casualties Percentage of total number of casualties 1960-1961 1961-1962 1960-1962 Cape Sparrow 28 30 10.9 Bokmakierie 30 13 81 Cape Weaver 12 27 73 Fiscal Shrike 18 20 71 Cape Turtle Dove 17 18 6.6 Cape Canary 14 20 64 Laughing Dove 20 12 6.0 Spotted Eagle Owl 12 8 3.7 Although the sequence within the first eight species diffe- red in the different years, the same eight headed the list of casualties in each of the two years. The sequence in Table 8 is based on the two years combined. DiscussION The number of bird road casualties recorded during the investigation no doubt only represents à part of the actual number. Badly squashed birds, especially when rather smal- lish or having been killed a long time ago are easily over- looked by the observer travelling in a car at a speed between 50 to 60 miles an hour. Moreover a considerable number of birds hit by a car will be thrown right off the road especially when hit by a fast travelling car. These will, therefore, not be noticed by an observer passing later. As has been shown the overall total casualty rate works out at 584 birds on just over 41500 miles. This is one bird to every 71 miles. It seems reasonable to assume that my own observations where the more accurate ones as during the two years period the investigation lasted 1 was always concentrating on spot- ting bird road casualties. If my records are separated from Source : MNHN. Paris Y'ORNITHOLOGIE PE A. Cape Sparrow, Passer melanurus & : B. Bokmakicrie, Telephorus seylonus : €. Laughing Dove, Streplopelia senegalensis ; D. Cape Turtle Dove, Streplopelia AT NE (BISL. OU | (Photographies G. + Broekhuysen) Source : MNHN. Paris L'OISEAU LA REVUE FRANÇAISE D'ORNITHOLOGIE Spotted Eagle Owl, Bubo africanus ; Cape Weaver, Ploceus capensis A, laris ; € canicollis. . Broekhuysen) (Photographies G. Source : MNHN. Paris BIRD CASUALTIES IN THE CAPE PROVINCE 39 the rest, it appears that during the first year I covered 8468 miles with a casualty rate of one bird per 48.1 miles. During the second year 10676 miles were covered and the casualty rate was one bird per 63.9 miles. If the two years are combined the casualty rate works out at one bird per 55.8 miles. This is a higher casualty rate than the one obtai- ned from the data tabulated in Table 1. Finnis (1960) summarises information on bird road casualties in Britain, France and America. He does not men- tion results obtained by J. McB. RoBerTson (1930) for Cali- fornia which work out at 63.9 miles per bird. MÔRZER Bruuns (1959) gives figures for Holland. The casualty rates expressed in number of miles per birds for the different countries are as follows : Britain France America Holland 1.5 142.5 9.8 9.9 43 563 63.9 6.3 1250 6.7 2444 12.5 16.6 42.7 61.2 In the case of the figures for Britain, in five instances the observer cycled, in two he walked and in one he was in a moving car. In all cases for France and America the observer was in a moving car while the figure for Holland was based on observations from a bicycle. Reinhold Hezn (1961) gives bird road casualties over a wide area in Germany but he does not give the actual casualty rate. From his data the casualty rate can only be assessed approximately. The figure works out at about 347 miles. BECKMANN (1961) counted the number of casualties along the same eight miles stretch every day for almost four years. This was also in Germany. He does not give the exact mileage covered during that time, but it can approximately be worked out and one then comes to a figure of one bird per c. 1220 miles. Both HEcp and BECKMANN observed from a car. The results obtained for the South Western Cape Province (one bird per 55.8-71 miles) is closest to the figures for Hol- land and Britain. It is clear that the number of birds killed in the South Source : MNHN. Paris 40 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Western Cape is much higher during the breeding-season than in the off-season. Reasons for the high casualty rate in the breeding-season are : — (a) the presence of juvenile and immature and, the- refore, inexperienced birds, (b) birds collecting building material for their nests and which due the activated buil- ding urge become less cautious, (c) birds collecting food for their chicks in the nest and probably paying less attention to traffic while collecting food on and near the road and while crossing the road on their way to the nest, (d) activated courtship and aggressive behaviour which gives rise to cha- sing behaviour, etc. The type of bird found as a casualty on the road varied tremendously, the two extremes being the Blue Crane on the one and the small Lesser Double-collared Sunbird, Crombek and White-eye on the other. The total number of 72 species identified seems rather high. If we total all the species men- tioned by Finnis (1956, 1960) and Hopson (1960, 1962) for British roads we come to 45. However, the avifauna of South Africa is considerably richer than that of Britain. There are different ways in which a bird can become a casualty and during the present investigation some of these ways became quite obvious. One would expect species which feed from the surface of the roads to become frequent casual- ties and Finnis (1956) mentions that members of the crow family are killed on the roads while feeding upon road casual- ties. The three members of the erow family in the Cape Pro- vince have the same habit and must be responsible for the removal of a considerable number of corpses of animals kil- led by traffic on the roads, but very few of them get run over themselves. In fact the only crow casualty among the 585 casualties was one Pied Crow. It is true that in this country crows are often shot at and are, therefore, extremely wary. Quite a number of birds of the lark and pipit families have the habit of sitting on the road, may be to dust-bath or because the road surface is warm. This probably accounts for the fairly high figure of 21 pipits and larks found dead on the roads. Birds of the weaver and seed-eater families are frequently seen on the road, feeding on droppings and in doing so may be run over by a speeding car. This is probably one of the reasons why the Cape Sparrow, Cape Weaver and Cape Source : MNHN. Paris BIRD CASUALTIES IN THE CAPE PROVINCE 41 Canary are among the eight species most frequently killed. This may to a certain extend also apply to the doves as the Cape Turtle Dove and the Laughing Dove are both fond of walking on the road, presumably while feeding but during the breeding-period also while courting. The high casualty rate among the Spotted Eagle Owls is probably also due to feeding on the road during the night (*). The blinding effect of head lights of an oncoming car will no doubt be an addi- tional lethal factor, in this case. Low flying and running birds are often killed whilst cros- sing the road. Good examples are the Bokmakierie, Fiscal Shrike, Boubou Shrike, Cape Thrush and some warblers which always fly low and usually in a straight line over the road. Francolins, especially during the breeding-season, often cross while leading their young from one side of the road to the other. If put to flight they move slowly and low over the ground. European Swallows often « hawk » low over tarred road surfaces and it is surprising that this species does not feature more promenently in the casualty figures. During a trip through the Eastern Cape Province in December 1961 I noticed many European Swallow casuallies on the road. Hesitation can be the cause of bird casualties. When cros- sing ahead of traffic, birds occasionally « jink » and take eva- sive action, sometimes even returning along their original course. Others fly off at a tangent but are overtaken and kil- led. In many cases these birds would have reached safety if they had continued along their original course or had flown up over the car. Most of the recorded casuallies were birds run over by other cars and only a small percentage concerns birds killed by the observer’s car. It is usually only in the latter cases that the reaction of the bird towards the approaching car can be observed. REACTIONS TOWARDS APPROACHING CAR Cape Sparrow (one observation by Miss. E. B. Robinson and three by G. J. Broekhuysen) : (a) Several flew on the road towards the car, all except one swerved (*) Hopsox (1962) mentions the habit of owls of swooping towards a moving light, especially during the breeding season in some country districts of Britain. 1 have never noticed this habit among South Afri- can species of owls. Source : MNHN. Paris 42 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE y just in time, but one dived right down in front of the car and killed, (@) One killed from a flock crossing the road in front of the car. () One 4 and one Q flew together crossing the road, they made no evasive movements and the & was hit and killed. The car moved at c. 58 miles an hour. () One bird while crossing the road, turned half way back when the car was noticed and escaped injury. Cape Francolin (two observations by R. K. Schmidt) : () Young francolins crossing the road crouched, spread all over ,the road and were difficult to avoid. One was killed. () Parent birds were trying to take their young across the road. Fiscal Shrike (two observations, one by G. J. Broekhuysen and one by R. K. Schmidt) : (a) An immature bird being fed on the road remained on the road although the parent bird flew off. The young bird was killed. () Bird flew up just in time and avoided collison with the car. Cape Turtle Dove (two observations, one by B. M. Mackenzie and one by R. Russell) : (a) Bird had been feeding in the gutter next to the road and was scared by passing car. @) Flew across the road and then noticing the approaching motoreyele turned towards the danger and actually hit the rider, but the bird was not hurt. European Swallow (two observations by G. J. Broekhuysen) (a) Bird was sitting on the road and did not get off in time. It was killed by a truck travelling at e. 45 miles an hour. () Bird flew fairly low over the road towards the approaching car and hit the roof. The bird could not be found although it was seen to fall on the road. Speckled Coly (one observation by R. K. Schmidt and one by Mrs. M. K. Rowan) (a) Bird crashed against windscreen of car travelling at 50-60 miles an hour. Bird showed no evasive action. () Flock started to fly across the straight open road, but turned back when danger was noticed. One bird was hit as it was turning. Bokmakierie (two observations by G. J. Broekhuysen) : Twice a Bokmakierie erossed the road in front of the car in a straight low flight and showed no evasive action. Cape Weaver (observation by R. K. Schmidt) : Bird flew straight against the car and there was no evasive action, The car was travelling at 60 miles an hour and the weaver was thrown about two yards off the road. Cape Thrush (observation by G. J. Broekhuysen) : Bird erossed the road flying low and straight just in front of the car and just missing it. No evasive action. Source : MNHN. Paris BIRD CASUALTIES IN THE CAPE PROVINCE 43 Cape Robin (observation by R. K. Schmidt) : Bird came out of the mist and was driven by a tailwind. When it noti- ced the car it showed evasise action but was hit at a speed of 25- 30 miles an hour. Cape Canary (observation by G. J. Broekhuysen) Bird flew straight down in front of the car. Feathers were flying but bird eould not be found. Cape Bunting (observation by C. J, Uys) When the road crossing bird saw the car it took successful evasive action, but then all of a sudden flew back on its previous course and was struck down. Orange-breasted Sunbird (observation by R. K. Schmidt) : Bird crossed the road and took no evasive action and was struck down at a speed of 25-30 miles an hour. APPENDIX ALPHABETICAL LIST OF THE SPECIES IDENTIFIED AS ROAD CASUALTIES African Quail Coturniæ coturniæ (L.) Bar-throated Apalis Apalis fhoracica (Shaw and Nodder) Blue Crane Tetrapteryx paradisea (Lichtenstein) Bokmakierie Telephorus zeylonus (L.) Boubou Shrike Laniarius ferrugineus (Gmelin) Bully Seed-eater Crithaga sulphurata (L.) Cape Bulbul Pyenonotus capensis (L.) Cape Bunting Fringillaria capensis (L.) Cape Canary Serinus canicollis (Swainson) Cape Dikkop Burrhinus capensis (Lichtenstein) Cape Francolin Francolinus capensis (Gmelin) Cape Longelaw Macromyx capensis (L.) Cape Robin Cossypha caffra (L.) Cape Sparrow Passer melanurus (Müller) Cape Thrush Turdus olivaceus L. Cape Turtle Dove Streptopelia capicola (Sundevall) Cape Wagtail Motacilla capensis L. Cape Weaver Ploceus capensis (L.) Cape White-eye Zosterops pallidus Swainson Cape Widow-Bird Coliuspasser capensis (L.) Cattle Egret Bubulcus ibis (L.) Chanting Goshawk Melierax musicus (Daudin) Common Tern Sterna hirundo L. Crombek Sylvietta rufescens (Vieillot) Crowned Cormorant Phalacrocoraz coronatus (Wahlberg) Crowned Guineafowl Numida meleagris (L.) European Starling Sturnus vulgaris L. European Swallow Hirundo rustiea L. Familiar Chat Cercomela familiaris (Stephens) Fiscal Flycatcher Sigelus silens (Shaw) Fiscal Shrike Lanius collaris L. Grassbird Sphenoeacus afer (Gmelin) Greater Honey-Guide Indicator indicator (Sparmann) Source : MNHN. Paris TABLE 1, — Bird casualties recorded on roads in the South Wes! vince during the period April 1960 - February 1962 and the mileage covered Species PHALACROCORACIDAE Crowned Cormorant Reed Cormorant ARDEIDAE Cattle Egret Yellow-billed Egret Egret sp. Night Heron ANATIDAE Duck sp. AQUILIDAE Steppe Buzzard Chanting Goshawk PHASIANIDAE Cape Francolin Grey-wing Francolin Red-wing Francolin African Quail NUMINIDAE Crowned Guinea-Fowl RALLIDAE Moorhen GRUIDAE Blue Crane CRARADRHDAE White-fronted Sandplover BurHINIDAr Cappe Dikkop LARIDAE Hartlaub’s Gull Common Tern CoLumsinar Cape Turtle Dove Laughing Dove Cape Turtle/Laughing Dove BUBONIDAE Spotted Eagle Owl CAPRIMULGIDAE S. A. Nightjar December 1960 | 1961 sm 1 ge EE RS: — Source - MNHN. Paris Species Gorupar Speckled Mousebird White-backed Mousebird Red-faced_ Mousebird Mouscbird sp. INDIGATORIDAE Greater Honey Guide ALAUDIDAE Thick-billed Lark Karroo Lark Red-capped Lark Long-billed Lark Grey-backed Finch Lark Lark sp. HIRUNDINIDAE European Swallow Larger Striped Swallow CORVIDAE Pied Crow PYGNONOTIDAE Cape Bulbul TURDIDAE Cape Thrush Mountain Chat Familiar Chat Sickle-wing Chat Stone Chat Cape Robin Karroo Scrub Robin SyLvDaE Crombek Grassbird Bar-throated Apalis Grey-backed_ Cisticola PRINIDAE Karroo Prinia MusGIGaPIDAE Fiscal Flycatcher January 1961 196: February| March d 1961 1964 1961] 1962 July November | August | September | October December 1961| 1960) 1961| 1960! 1961 | 1960 | 1961 | 1960] 1961 | 1960 | 1961! 1960! 1961 1 1 1 1 1 1 1 L et 1 1 1 1 2 | 1 1 æ 1 3 1 1 1 1 1 1 1 1 1 1 1 are SE 2 1 1 | 1 1 à | 1 À 1 “ni | 1 | 1 1 l De: 1 4 Source - MNHN. Paris Species January | Fébruary| March | April june July August (September | October | November | December 1961, 1962 1961, 1962 1961, 1962 1960 1961, : | 1961 1960 |1961 11960 11961 11960 1961 1960 1961 |1960 |1961 |1960 1961 MOTACILLIDAE | Ve)! Cape Wagtail ñ Pipit sp. 1 | Pipit or Lark Cape Longelaw LANTIDAE Fiscal Shrike n4 PAIE ES eIEt Boubou Shrike 1 Bokmakierie NAT 1 5 ET STuRNIDAE European Starling Wattled Starling Pied Starling 1 1 Red-winged Starling 1 | PROMEROPIDAE | Sugarbird 2 NECTARINIDAE Orange breasted Sunbird RU Lesser Double-collared Sunbird | ZOSTEROPIDAE | Cape White-eye à | PLOGEIDAE Cape Sparrow APE Cape Weaver ans Red Bishop Bird 1 Cape Widow-bird Tu 0 QU 1 Red/Cape Widow-bird | el w » FRINGILLIDAE | Cape Canary 3 | Bully Seed-eater 1 ï Yellow Canary | White-throated Seed-eater el Seed-cater sp. Ke: 1 1 Cape Bunting Unidentified MRC RSA oite 1 Pa a Fr 9 1 + | 2 19 12 11 13 21 15 39 29 69 21 127 15 #4 100 Month total in number 45 | 14 | 18 11! 25 | 12 | 12 | 16 10: | 9 13 14 un os ir |25 ao |at | 10 | 2 | 16 Number of different species 25 2 1e: 6 | 14 7 9 Ki MG Lnsan jaça7 Laane |2700 laure Joss |oo1a 1aas Lisaë 1181 | 2206 | 1852 ! Û Monthly mileage 12629 1282/1264; 947)2266|1031|1249) 2038; Source - MNHN. Paris 50 L'OISEAU ET LA REVUE Grey-backed Cisticola Grey-backed Finch-Lark Grey-wing Francolin Hartlaub’s Gull Karroo Lark Karroo Prinia Karroo Serub Robin Larger Striped Swallow Laughing Dove Lesser Double-collared Sunbird Long-billed Lark Moorhen Mountain Chat Night Heron Orange-breasted Sunbird Pied Crow Pied Starling Red Bishop Bird Red-capped Lark Red-faced_Mousebird Redwing Francolin Red-winged Starling Reed Cormorant Sickle-wing Chat South African Nightjar Speckled Mouscbird Spotted Eagle Owl Steppe Buzzard Stone Chat Sugarbird Thick-billed Lark Wattled Starling White-backed Mousebird White-fronted_Sandplover White-throated_Seed-eater Yellow-billed Egret Yellow Canary FRANÇAISE D'ORNITHOLOGIE Cisticola subruficapilla (A. Smith) Eremopterix verticalis (A. Smith) Francolinus africanus Stephens Larus hartlaubii (Bruch) Certilauda albescens (Lafresnaye) Prinia maculosa (Boddaert) Erythropygia corgphaens (Lesson) Cecropsis eucullata (Boddaert) Stigmatopelia senegalensis (L.) Cinnyris chalybeus (L.) Certhilauda curvirostris (Hermann) Gallinula chloropus (L.) Oenanthe monticola Vieillot Nycticorax nycticoraæ (L.) Anthobaphes violacea (L.) Corvus albus Müller Spreo bicolor (Gmelin) Euplectes orix (L.) Tephrocorgs cinerea (Gmelin) Urocolius indicus (Latham) Francolinus levaillanti (Valenciennes) Onychognathus morio (L.) Phalacrocoraz africanus (Gmelin) Cercomela sinuata (Sundevall) Caprimulgus pectoralis Cuvier Colius striatus Gmelin Bubo africanus (Temminck) Buteo buteo (L.) Saxicola torquata (L.) Promerops cafer (L.) Calendula magnirostrus (Stephens) Creatophora cinerea (Menschen) Colius colins (L) Charadrius marginatus Vieillot Crithagra albigularis A. Smith Mesophoyx intermedius (Wagler) Crithagra flaviventris (Swainson) RESUME 1) En deux années de recherches dans la partie Sud-Ouest de la Province du Cap (Afrique du Sud), l’auteur, après avoir parcouru 41500 milles (74700 km), a dénombré 584 oiseaux tués par accidents de la route. 2) 72 espèces furent identifiées. 3) Le taux d’accident pendant la période de reproduction s'avère à peu près le double de celui hors de la saison de nidification. 4) Le taux d’accident sur les routes de la zone étudiée est comparé aux données recueillies en Grande-Bretagne, France, Allemagne et Amérique. Source : MNHN. Paris BIRD CASUALTIES IN THE CAPE PROVINCE 51 5) Les différents modes possibles d'accidents sont étu- diés ; quelques observations sur le comportement des oiseaux traversant la route devant un véhicule sont rapportées. REFERENCES Barnes, M. D. (1936). — The death-toll of birds on our ronds. Natura- dist : 89. BeCKMANX, H. (1961). — Vogelverluste auf Autostrassen. Orn. Mitt., 13 :128. Finnis, R. G. (1956). — Bird casualties on roads. B.0.C., 76 : 150. Finnis, R. G. (1960). — Road casualties amongst birds. Bird Stud., 7 : 21. Heun, R. (1961). — Vogelverluste auf Autostrassen. Orn. Mitt., 13 : 202. Hopsox, N. L. (1960). — A survey of vertebrate road mortality 1959. Bird Stud., 7 : 224. Hopsow, N. L. (1962). — Some notes on the causes of bird road casual- ties. Bird Stud, 9 : 168. LinspaLe, J. M. (1929). — Roadways as they affect bird life. Condor, 31: 143. Mantexs, J. (1962). — Gefahrung der Vogelwelt durch Kraftwagen. Orn. Mitt., 14 : 221. Môrzen Bauuns, M. F. (1959). — Fauna sterfte door het verkeer. Leo. Nat. 62: 73-717. Rogerrson, McB. J. (1930). — Roads and birds. Condor, 32 : 142. Scnoncen, A. W. (1954). — A study of roadkills. Passenger Pigeon, 16:56 Waure, F, B. (1927). — Birds and motor cars. Auk, 44 : 265. Waure, F, B. (1929). — Birds and motor cars. Auk, 46 : 377. Source : MNHN. Paris LES PIGEONS DE LA NOUVELLE-CALEDONIE par J. DELACOUR Secrétaire général du IX° Congrès International d'Ornithologie (Rouen 1938) Le goût des oiseaux et leur étude a eu tout d’abord, pour Jacques BERLIOZ, comme pour moi, une base esthétique. Dès notre plus jeune âge, nous les avons aimés parce qu’ils sont beaux. Les aimant, nous avons voulu les connaître, et cela décida de nos carrières. Bientôt nous avons dû envisager les problèmes nombreux qu'ils présentent, et les multiples aspects qu'ils revêtent. Cela fait peu à peu oublier le véri- table point de départ. La beauté des oiseaux est présente dans presque tous les différents ordres ; parmi tant d’autres, deux de ces derniers sont particulièrement attrayants par la singularité des carac- tères, par le nombre et variété des genres et des espèces dont le plumage, toujours élégant, est souvent même d’un éclat extraordinaire. Leurs formes, leurs mœurs très spéciales, attirent l'attention. Je veux parler des Perroquets et des Pigeons, qui sont abondamment répandus dans toutes les parties du monde où le climat leur est favorable. Le plus grand nombre d’entre eux habitent les tropiques, mais on en trouve aussi dans des régions tempérées plus ou moins froides en hiver. Ces deux ordres d'oiseaux ont en outre l’avantage d'être représentés largement dans les îles, même celles de dimensions restreintes, particulièrement en Océanie, où des espèces somptueusement ornées existent un peu partout avec une exhubérance de variation vraiment étonnante. J'ai eu l'occasion, au cours des trois dernières années, de visiler à deux reprises l’une des plus vastes et des plus riches, au point de vue de la faune avienne, des îles du Pacifique Méridional : la Nouvelle-Calédonie. J'ai pu y observer dans la nature la plupart des quelque 100 espèces qui l’habitent. J'ai aussi constaté avec beaucoup de satisfaction l'intérêt que portent à l'ornithologie un bon nombre de ses habitants ; à tel point que je me suis décidé à publier à brève échéance L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris PIGEONS DE LA NOUVELLE-CALÉDONIE 53 un manuel des oiseaux calédoniens qui permettra à ceux qui les rencontrent de les identifier et d'apprendre l'essentiel de ce qui les concerne. Les espèces de Perroquets et de Pigeons y sont singulière- ment nombreuses pour l'étendue du territoire. Il en existe quatre des premiers, et Jacques BERLIOZ en a publié une excellente étude (L’Oiseau et la R.F.0., XV, N.S., 1945, P. 1). Quant aux seconds, je vais essayer de les évoquer ici. La Région Calédonienne, composée de la Grande Terre, des îles voisines, et des trois Loyauté : Maré, Lifou et Ouvéa, ne possède pas moins de six espèces autochtones, auxquelles s’en est ajoutée une autre, introduite du sud-est et de l'Asie : Streptopelia trigrina. Cette dernière s’est parfaitement accli- matée ; elle vit aux alentours des habitations humaines, dans les jardins et les cultures et, de ce fait, elle ne nuit aucune- ment aux espèces indigènes, qui ne fréquentent que les forêts ou les savanes boisées. Des six espèces locales, l’une forme un genre séparé ; une seconde est endémique ; une troisième occupe aussi les Nou- velles-Hébrides ; une quatrième est représentée par une sous- espèce particulière, landis que les deux autres appartiennent à des sous-espèces existant aussi, l’une dans l’ouest du Paci- fique (des Salomons à Samoa), l’autre en Australie. Quatre des pigeons calédoniens appartiennent au groupe des Carpophages, les Tréroninés ; les deux autres à celui des Columbinés granivores. Les affinités de ces espèces sont diverses. La première, Ptilinopus greyi, ne se rencontre ailleurs qu'aux Nouvelles- Hébrides, aux Iles Banks et Santa Cruz ; elle est voisine de P. porphyraceus qu’on trouve, sous plusieurs formes, à Samoa et aux Fijis. La seconde, Drepanoptila holosericea, est complè- tement distincte et ne paraît pas avoir de proches parents. La troisième, Ducula pacifica, vit aussi dans de nombreuses iles, depuis celles au large du nord-est de la Nouvelle-Guinée, Ellice et Phoenix, jusqu’à Samoa, les Fijis et Tonga. La qua- trième, Ducula goliath, l'énorme « Notou », est endémique, et ses seuls cousins, assez éloignés, sont D. brenchleyi, des Salomons, D. bakeri, des Nouvelles-Hébrides, et D. latrans, des Fijis. La cinquième, Columba vitiensis, est une espèce largement répandue des Philippines et des Petites Iles de la Sonde à Samoa, sous des formes variées, et elle est apparentée à C. janthina, du Japon. Tous ces pigeons sont donc d’aff- Source : MNHN. Paris 54 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE nités nettement océaniques. La sixième, Chalcophaps indica, est sans doute d'origine australienne puisqu'elle est repré- sentée en Nouvelle-Calédonie par la sous-espèce chrysochlora de ce continent. L’aire de dispersion de l'espèce comprend toute la région Indo-Malaise, de l'Inde aux Philippines, aux Moluques, à l'Ile Christmas, aux archipels d’'Entrecasteaux et de Louisiade, et à la Nouvelle-Guinée, de la Baie de l'As- trolabe, au nord, au Hall Sound, au sud. Voici la liste des pigeons de la Région Calédonienne : PTILOPE DE GREY (Ptilinopus greyi) Petit (20 cm). Couronne rouge violacé ; reste de la tête, cou, poitrine et côtés gris-vert clair ; manteau, ailes el queue verts, cette dernière avec bande terminale grise; milieu du ventre rouge carmin ; sous-caudales orange ; œil brun ; bec gris-vert ; pattes rouges. La femelle est un peu plus terne. Les jeunes n'ont pas de rouge, et le dessous du corps est vert légèrement strié de jaune. Cette jolie Colombe se nourrit surtout de petites figues de banyan et de baies. Elle est répandue dans les bois et les ver- gers, se déplaçant à la recherche de sa nourriture. Elle habite les Iles Loyauté, les Nouvelles-Hébrides et îles voisines, mais elle est peu commune en Nouvelle-Calédonie. Après les oura- gans, on la trouve en grand nombre sur la grande terre, par- ticulièrement dans le sud et à l'Ile de Pins, le vent violent l'ayant emportée des Loyauté. Elle diminue en nombre par la suite et, bien qu'elle niche, elle demeure rare jusqu’à ce qu'un nouveau contingent soit amené par une autre tempête. PIGEON VERT SOYEUX (Drepanoptila holosericea) Assez grand (mâle : 28 em). Tête, cou, poitrine et parties supérieures vert brillant, avec une mince bande blanche du menton à la gorge, où elle se termine en pointe, et des bandes gris perle aux ailes et à la queue, qui est très ample ; les plumes du front et de l'avant de la couronne sont allongées et serrées formant une sorte de huppe touffue au-dessus du bee, qui est court et noir verdâtre ; une double barre étroite, Source : MNHN. Paris PIGEONS DE LA NOUVELLE-CALÉDONIE 55 blanche et noire, au bas de la poitrine ; ventre jaune ver- dâtre ; bas-ventre et dessous de la queue jaune d'or, les sous- caudales aussi longues que les rectrices ; deux grosses touffes blanches sur les pattes, qui sont rouge carmin et presque cachées ; œil brun-rouge. Les rémiges sont divisées à l’extré- mité, les deux moitiés incurvées vers l'extérieur, caractère tout à fait unique. Femelle bien plus petite (23 em) et plus terne, mais montrant, atténuées, toutes les marques du mâle ; les jeunes lui ressemblent de couleur. Cette superbe espèce constitue un genre spécial, propre à la Nouvelle-Calédonie ; c’est sans doute le plus bel oiseau de la région, et, avec le Kagou, le plus intéressant et le plus particulier. Elle habite les forêts et les bois des savanes, et elle est souvent difficile à voir, car elle se tient immobile et se confond avec le feuillage, ne se signalant que par son fort et rauque roucoulement. Elle jouit en principe d'une pro- tection totale, et il faut espérer que tous respecteront désor- mais ce joyau de l’avifaune calédonienne. Elle a malheureu- sement été décimée par une chasse abusive dans le passé. CARPOPHAGE PACIFIQUE (Ducula pacifica pacifica) Grand (40 cm). Bec noir, faible, surmonté à la base d’une caroncule noire liserée de blanc, plus développée chez le mâle. Tête, cou et haut du dos gris clair ; dessous du corps gris vineux ; sous-caudales marron; dos, manteau, ailes et queue vert-bleu, foncé et métallique ; pattes rouges. Femelle légèrement plus petite et plus terne. Cette belle espèce se rencontre dans l’ouest du Pacifique. Elle est commune aux Loyauté, mais ne paraît visiter qu'irré- gulièrement la Nouvelle-Calédonie où, comme le Plilope de Grey, elle est amenée périodiquement par les ouragans. Elle niche et s'établit, mais diminue en nombre assez rapidement. Se réunit en bandes, volant d’ile en île à la recherche des fruits dont elle se nourrit. Roucoulement profond et rauque. CARPOPHAGE GOLIATH (Ducula goliath) Très grand (50 em). Gris ardoisé noirâtre, avec du marron aux ailes, au ventre et à la queue, et du blanc aux parties Source : MNHN. Paris 56 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE anales ; œil orange ; bec et pattes rouge carmin. Appelé loca- lement Notou. C’est l’un des oiseaux les plus particuliers et les plus inté- ressants de la Nouvelle-Calédonie, et aussi le plus gros de tous les Pigeons arboricoles du monde, Seuls les Gouras de la Nouvelle-Guinée, espèces terrestres, le dépassent en taille. Ce splendide Carpophage fréquente les forêts, où on entend résonner sa voix tonnante et étrange. Il se nourrit de fruits. 11 a malheureusement été décimé par une chasse irraisonnée, et il est urgent de le protéger par des mesures efficaces, telles qu'une très courte saison d'ouverture, et surtout la création de réserves. PIGEON À GORGE BLANCHE (Columba vitiensis hypoenochroa) Grand (40 em). Tête, cou, haut du dos et dessous du corps marron pourpré brillant, avec la gorge et les joues blanches ; reste du dessus du corps gris foncé avec de riches reflets verts sur le dos ; bec, paupières et pattes rouges. Jeunes plus ternes, avec la gorge grise. C’est un véritable Pigeon, avec un bec plus dur que celui des espèces précédentes. Il habite toute la Région Calédo- nienne. Il se déplace dans les forêts et les savanes, souvent en troupes, à la recherche des baïes et des graines, effec- tuant des voyages parfois considérables. Il niche assez bas, sur les pandanus, souvent même sur le sol, en particulier sur la grande terre. Cette belle espèce est trop persécutée et diminue en nombre. Il est indispensable de réglementer plus sérieuse- ment sa chasse et de la protéger. CoLoMBE TURVERT (Chalcophaps indica chrysochlora) Assez petite (25 em). Queue courte et pattes hautes. Tête, cou, parties inférieures et manteau roux vineux, plus pâle en dessous ; ailes vert émeraude métallique avec une petite tache gris pâle à l'épaule et les rémiges brun foncé ; dos noir avec deux bandes grises ; queue brun foncé ; œil brun ; bec orange ; pattes rouges. Source : MNHN. Paris PIGEONS DE LA NOUVELLE-CALÉDONIE 57 Cette belle Colombe habite la Région Calédonienne, les Nouvelles-Hébrides et l'Australie ; d’autres sous-espèces, la plupart avec la couronne grise chez le mâle, occupent le sud de l’Asie, la Malaisie et les régions situées au sud-est. C'est une espèce terrestre, qui se nourrit de graines tombées. Elle vit sous bois et dans les savanes, se cachant dans les arbres épais, et elle n’est pas facile à voir, bien qu’assez commune ; on la surprend parfois traversant au vol un sentier ou une clairière. Ce vol est rapide et direct, et elle se perche fréquemment. Niche sur les arbres bas. Source : MNHN. Paris QUELQUES PARTICULARITES DE LA REPRODUCTION DES OISEAUX DANS LES HAUTES ZONES DU THIAN-CHAN par G. P. DEMENTIEV et L. S. STÉPANYAN Musée Zoologique de l'Université (Moscou) Les renseignements recueillis dans les hautes montagnes de Thian-Chan, au cours d’expéditions et d’excursions effec- tuées par L. STépanvaN pendant la saison de reproduction des années 1953, 1954, 1956, 1957, 1959, 1960, ont servi de base à cet article. Les données indispensables pour la comparaison (nombre de pontes, quantité d'œufs pondus chez les populations de plaines de certaines espèces largement répandues) sont tirées de publications diverses, judicieusement condensées dans la Monographie « Les oiseaux de l’Union Soviétique ». En revanche, les données sur le taux de fécondité des oiseaux en haute montagne sont toutes originales, vu que dans les ouvrages spécialisés, à de rares exceptions près, l’altitude pré- cise des lieux de nidification n’est pas indiquée, alors que ce fait présente une importance décisive lors de la comparai- son de la fécondité des populations d’une même espèce en fonction de leur répartition verticale. Dans cet article, les étages subalpins et alpins ont été réunis sous le terme de < haute montagne » el se situent à une altitude de 2900 à 4000 m (et parfois même plus) au-dessus du niveau de la mer. L'écologie des populations d’une espèce donnée est à la base des problèmes touchant l'écologie de l'espèce dans son ensemble. Néanmoins, les méthodes actuelles de recherche ne permettent pas toujours de détecter ni d’éclaircir d’une façon satisfaisante les particularités de l’existence de l'espèce dans les différentes parties de sa zone de répartition. Il nous arrive souvent d’avoir affaire, pour une même espèce, à toute une série de formes géographiques morphologiquement bien caractérisées pour lesquelles les différences écologiques ne L'Oiseau et R.F.O0. V. 35, 1965, n° spécial. Source : MNHN. Paris REPRODUCTION DES OISEAUX DU THIAN-CHAN 59 sont pas toujours suffisamment nettes. Souvent les races d’une même espèce ne diffèrent que par des détails de biologie relativement insignifiants. Ainsi, elles ont par exemple dés dates variables pour le début et la fin de certains cycles vitaux : reproduction, mue, etc. De telles différences sont le plus souvent observées chez les espèces largement répandues, mais néanmoins, dans les limites de l'aire de répartition où les conditions écoclima- tiques conservent leurs caractères principaux. Dans ces zones les parties optimales et minimales sont mal déter- minées. Par ailleurs, lorsque l’espèce possède une haute plas- ticité écologique elle peuple des étendues dont les caractéris- tiques de certaines parties diffèrent sensiblement. Dans les limites de l'aire de distribution d’une telle espèce les zones les plus comme les moins favorables sont nettement exprimées. Les différences écologiques des formes géogra- phiques de telles espèces eurytopes ont un caractère sensi- blement plus profond. On observe chez elles, en plus des particularités phénologiques, des différences permanentes dans l'intensité et les rythmes de reproduction. Comme on le sait, la fécondité et la mortalité sont toujours en stricte corrélation entre elles et, dans les conditions normales, la majorité des espèces animales a, quantitativement, une limite d'oscillation relativement peu sensible. Cependant, comme l'ont démontré les résultats des recherches de certains auteurs, généralisés et analysés par D. Lack (1954), la fécon- dilé est un phénomène par lui-même labile de l'organisme et varie largement selon les conditions, même chez un seul individu. Néanmoins elle a, dans tous les cas, un caractère excessivement adaplatif. Et, comme les recherches précitées l'ont démontré, le nombre d'œufs de la ponte correspond tou- jours aux possibilités d'élever la quantité maximum d'’indi- vidus normaux de la jeune génération. L'analyse de ce phénomène dans les milieux désertiques d'Asie Centrale (qui peuvent être considérées comme zones défavorables pour toute une série d'espèces) a été faite par A. K. RousrAMOv (1954). Les hautes montagnes représentent (semble-t-il) une zone défavorable pour de nombreuses espèces largement répandues. Un des facteurs probables qui réduit le nombre de ces espèces paraît être la limitation des réserves de nourriture par les Source : MNHN. Paris 60 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE conditions rigoureuses de ces lieux. Cette dépendance se mani- feste particulièrement chez les oiseaux insectivores. L’inten- sité de leur reproduction est ici nettement moins importante que dans les populations de plaines. Il est à noter que la majorité des espèces nidifiant en haute montagne n'ont habituellement qu'une ponte par an. Sur les hautes mon- tagnes aussi bien que dans d’autres parties de l’aire de répar- tition, des pontes de remplacement ont également lieu. Cepen- dant, aux grandes altitudes, ce phénomène se rencontre plus rarement, vu la durée beaucoup plus brève de la saison de reproduction. Eremophila alpestris albigula Bp. est la seule espèce dont nous ayons observé un cycle reproductif normal. Sur les hautes montagnes du Thian-Chan (3500 à 4000 m d'altitude), dans la deuxième quinzaine de juillet 1956, eut lieu la sortie en masse des petits de ces oiseaux. Des adultes se sont remis à pondre tout en continuant à nourrir leurs jeunes. Toutefois nous constatâmes que ceci ne concernait pas tous les couples, car une partie d’entre eux se contenta d’une seule ponte. Néanmoins, le fait que la répétition normale du cycle reproductif en haute montagne n’affecta qu’une partie de la population, mérite incontestablement notre attention. Pour comparer le nombre d'œufs dans la ponte des popu- lations de plaine et de haute montagne, nous avons utilisé une série d’espèces largement répandues formant dans les montagnes des sous-espèces nettement caractérisées (voir tableau). Cependant, il est à noter que toutes les populations des sous-espèces montagnardes ne sont pas caractérisées par une fécondité plus basse. Ceci n’est vraisemblablement propre qu'aux groupes d'individus qui peuplent précisément les hautes montagnes mais non pas les montagnes en général. Dans ce cas, les limites écologiques et morphologiques de l'isomorphisme géographique peuvent ne pas correspondre entièrement. Dans le tableau ci-dessous nous n’avons utilisé que les données concernant les nids trouvés dans les limites des zones alpines et subalpines, à l'exception de Motacilla citreola Pall. et M. flava L., dont les nids ont été trouvés à une alti- tude de 2400 m. Ainsi, d'après le tableau ci-dessous, malgré une certaine insuffisance des données, on observe facilement une tendance à la réduction du nombre d'œufs par ponte chez les popu- Source : MNHN. Paris REPRODUCTION DES OISEAUX DU THIAN-CHAN NOMBRE D'ŒUFS PAR PONTE Espèces et sous-espèces Phylloscopus inornatus inornatus Phylloscopus inornatus humei Motacilla citreola werae et Motacilla flava flava Motacilla citreola calcarata et Motacilla flava feldegg Anthus _trivialis trivialis Anthus trivialis haringtoni Phoenicurus phoenicurus Phoenicurus aurorens Phoenicurus coeruleocephalus (1) Phoenicurus ergthronotus Phoenicurus erytrogaster (1) La littérature donne pour la ponte de Ph. Nombre de nids examinés ou source de renseignement Renseignements puisés dans la littérature Renseignements puisés dans la littérature Renseignements puisés dans la littérature Renseignements puisés dans la littérature Renseignements puisés dans la littérature 2 5 10 (renseignements puisés dans la littéra- ture) 2 6 (renseignements puisés dans la littéra- ture) 3 à 5, soit en moyenne 4 œufs. 61 Nombre d'œufs dans la ponte max, 6 Û & min, 5 3 3 moy. 5 4,2 6,5 4,1 coeruleocephalus de Source : MNHN. Paris 62 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Jations de haute montagne des espèces largement répandues en comparaison de celles des plaines. Il serait très intéressant de faire sur ce plan une comparaison entre certaines espèces spécialisées de montagne el de plaine rapprochées entre elles du point de vue systématique. Nos données permettent d’ef- fectuer une telle comparaison pour les espèces du genre Phoenicurus. Les données sur les espèces de montagne sont originales ; celles concernant les espèces de plaine sont tirées de la littérature scientifique. Ph. phoenicurus L. habite l’Europe et les parties centrales de l'Asie jusqu'au lac Baïkal et les monts Sayan-Oriental, à l'est. A l'exception des populations méridionales, la plus grande partie de l'aire de répartition se situe dans les plaines. Ph. auroreus Pall. habite les régions autour du lac Baïkal jusqu'aux monts Kentey en Mongolie. La plus grande partie de l'aire de répartition embrasse la Mandchourie orientale, la Corée, la Chine à partir des régions du nord-est jusqu’au Yunnan. L'oiseau se trouve de préférence dans les forêts de plaine, au pied des montagnes. Ph. coeruleocephalus Vig. vit sur les montagnes de l'Asie centrale, de l'Afghanistan, du Beloutchistan, de l'Himalaya, à l'est jusqu’au Sikkim et au Boutan. Biotope : forêts de conifères de montagne. Ph. erythronotus Ev. vit sur les monts de l’Asie centrale, à l’est jusqu’au Baïkal méridional, Khangaï, Tarbagatai. Bio- lope : les bois de montagne et les buissons ; en Asie centrale, la partie supérieure de la zone des forêts de conifères. Ph. erythrogaster Güld. peuple le Caucase, les monts de l'Asie centrale, l'Altaï et les régions situées au nord du Baï- kal. Partout cet oiseau de haute montagne est étroitement lié aux conditions alpines. Les données ci-dessus mentionnées peuvent être com- plétées par certains renseignements concernant principale- ment le nombre de petits par couvée qui attestent une fécon- dité inférieure de la sous-espèce de Parus alricapillus songarus Sev. en comparaison avec P. a. borealis Sel. ; ainsi que des populations de haute montagne de Turdus viscivorus bonapartei Cab. comparée au T. ». viscivorus L. européen. D'après les données tirées des ouvrages spécialisés, le nombre d'œufs dans une ponte de Luscinia calliope calliope Pall. de Source : MNHN. Paris REPRODUCTION DES OISEAUX DU THIAN-CHAN 63 plaine est de 4 à 6 alors que chez l'espèce de haute montagne L. pectoralis Gould elle va de 3 à 5 œufs. Enfin, il est intéressant sous ce rapport de faire une com- paraison entre les espèces de haute montagne et de plaine, qui du point de vue écologique ont beaucoup de points com- muns, mais phylogénétiquement sont loin l’une de l’autre. Tels sont, à notre avis Regulus regulus L. et Leptopoecile sophiae Sev. Les dimensions absolues du corps, la nourri- ture, la nidification et dans une certaine mesure le biotope de ces espèces ont beaucoup de points communs. Une ponte normale de R. regulus en Europe se compose de 8 à 9 œufs. La ponte normale de L. sophiae dans la zone subalpine de Thian-Chan ne dépasse probablement pas 4 à 5 œufs. On peut ainsi supposer que la tendance vers la réduction de la fécondité sur les hautes montagnes est propre à plu- sieurs espèces d'oiseaux. Il est hors de doute que lors des recherches ultérieures toute une série de telles espèces sera révélée. Il est indispensable de souligner une fois de plus que ce phénomène est principalement propre aux oiseaux insecti- vores. Parmi les causes des faits examinés, le caractère des sources nutritives dans les hautes montagnes a une impor- tance primordiale. Il n'est pas exclu qu’un certain rôle puisse être joué ici par le type de la balance énergétique dans les conditions du climat rigoureux de haule montagne. Néanmoins, il faut par- ticulièrement souligner combien les tentatives actuelles d’ex- plication du phénomène examiné ont un caractère hypothé- tique, car cette question exige une étude spéciale et de plus amples données. En tout cas, les hautes montagnes du Thian- Chan central possèdent une faune entomologique beaucoup plus pauvre, quantitativement et qualitativement, que les zones siluées plus bas. Cependant, durant la période de nidi- fication, toules les espèces et sous-espèces d'oiseaux de haute montagne ci-dessus mentionnées se nourrissent, elles-mêmes el leurs petits, d'insectes. Ceci est démontré par la dissection des organes digestifs et par l'observation directe. Par exemple Ph. erythrogastler, qui se nourrit principale- ment de plantes, alimente, sur les plateaux, ses petits d’in- sectes et surtout de chenilles trouvées dans les herbes des- séchées qui constituent la flore locale. Les araignées, large- ment répandues sur les hautes montagnes, jouent un grand 5 Source : MNHN. Paris 64 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE rôle dans la nourriture des insectivores de ces lieux, cepen- dant elles ne suffisent pas à constituer complètement la base alimentaire de ces espèces et cette dernière reste évidemment plus pauvre que celle, par exemple, au pied des monts ou dans la partie inférieure de la zone forestière. Il est possible que l’insignifiante quantité générale de tous les oiseaux insec- tivores de haute montagne soit liée à ce facteur limitatif. L'action permanente de ce facteur a provoqué l'apparition de certains caractères adaptatifs dans l’écologie des espèces avec une spécialisation alimentaire correspondante. Une fécondité basse comme particularité des populations de haute montagne d'espèces largement répandues, semble donc un caractère nettement adaptatif et doit être envisagé comme l'effet phy- siologique des conditions d'existence. En cela se concrétise également une des manifestations de l'isomorphisme géographique sous son aspect écologique. Source : MNHN. Paris SUR LE FAISAN DE HUME ET LE GENRE CALOPHASIS par Alessandro Gnigr Ancien Recteur de l'Université de Bologne Président du IX*° Congrès International d'Ornithologie (Rouen 1938) Le Faison de Hume fut découvert en 1881 dans le Manipur par HUuME lui-même, mais il n’eut pas la possibilité de mener des recherches personnelles sur cette espèce. En effet, le seul couple qu’il possédait lui avait été donné par des indigènes, desquels il avait appris que ces Faisans vivaient dans des forêts épaisses à une altitude oscillant entre 800 et 1500 m ; ils n'étaient ni fréquents, ni abondants, dans la région qui s'étend jusqu'à la Birmanie Nord-Ouest. Hume voulu dédier à sa femme cette espèce nouvelle, mais au lieu d'écrire, comme il eût été correct, humei, il féminisa son propre nom d’une façon discutable puis le mit au géni- tif, d’où : humiae. D'après DELACOUR, peu d’exemplaires furent capturés, par la suite peu de résultats satisfaisants furent obtenus en ce qui concerne l'élevage en captivité, enfin aucune importation en Amérique ou en Europe n'eut lieu avant la publication de son grand ouvrage : « The Pheasants of the World » paru en 1951. En février 1963, M. George Munno, de Calcutta, m'offrit des Faisans de cette espèce. J'acquis deux mâles et une femelle. Le deuxième mâle fut accouplé avec une femelle d'Elliot. En 1963 et en 1964 j'obtins des deux couples de nombreux jeunes très forts que j'ai élevés très facilement. Ils m'ont ainsi permis de faire un second croisement corres- pondant à la formule humiae X (humiae X ellioti). NOTES DESCRIPTIVES SUR Calophasis humiae HUME Je ne donnerai pas ici une description de l'espèce, mais je voudrais souligner les différences saillantes entre C. Aumiae et C. ellioti. L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris 66 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Le mâle de humiae est légèrement plus pelit et plus agile que le mâle d’ellioti. Ils ont, tous les deux, la même voix et la même démarche. La queue de humiae est plus longue, elle atteint en effet 400-535 mm de long, tandis que celle de ellioti oscille entre 390 et 440 mm. La longueur totale moyenne est de 900 mm pour humiae, de 800 mm pour ellioti, aussi la première espèce paraît-elle plus élégante et plus élancée que la seconde. En ce qui concerne les couleurs du mâle, le caractère le plus frappant est la couleur du cou, entièrement métallique, changeant du vert à l’acier, aussi bien dans la région dorsale (brun blanchâtre chez ellioti) que sur les côtés (blancs chez ce dernier). En outre chez ellioti les couleurs du cou se déta- chent très nettement des teintes rouge cuivre du dos et de la poitrine qui tranchent sur le blanc du ventre, tandis que chez humiae les nuances vertes passent graduellement au châ- tain foncé el brillant pour gagner toutes les parties infé- rieures. Les parties supérieures, dos et ailes, sont châtain foncé chez humiae et rouge cuivre chez ellioti. Le dessin général de l'aile est le même chez les deux espèces mais on peut remarquer des différences entre les trois bandes respectivement blanches et noires métalliques des ailes. La bande blanche des scapulaires est moins évidente chez humiae, tandis que celle qui traverse les couvertures alaires est plus large chez humiae que chez ellioti. La tache elliptique placée au milieu de l'aile, noir luisant avec une légère nuance métallique chez ellioti, est plus ample et changeante (du vert à l’acier) chez humiae. Les dessins du croupion et de la queue sont différents chez les deux espèces : chez ellioti le dos et le croupion sont à bandes alternées blanches et noires, chez humiae les bandes blanches alternent avec des bandes bleu acier ; la queue de l'Elliot montre une alternance de larges bandes grises et de bandes châtain séparées les unes des autres par des bandes noires irrégu- lières ; sur la queue de humiae, de larges bandes grises sont alternées de bandes noires terminées par une mince bande brune. La différence principale entre les femelles demeure dans la gorge, qui chez humiae est d’un gris pâle immaculé, tandis Source : MNHN. Paris FAISAN DE HUME ET GENRE CALOPHASIS 67 que chez ellioti elle présente une tache noire en forme de bavette qui descend jusqu’à la base du cou. DISTRIBUTION GÉOGRAPHIQUE DU GENRE Calophasis Si l’on commence par l'Occident, on trouve C. humiae dans le Manipur et l’Assam, en contact avec C. burmanicus de la Birmanie dont la silhouette rappelle C. ellioti avec la tête et le cou de C. humiae. À une distance considérable on trouve en Chine orientale, au sud du Fleuve Yang-tsé-Kiang, C. ellioti et au sud-est, dans l’Ile de Formose, C. mikado. HYBRIDATION €. mikado fut découvert au début de ce siècle dans l’Ile de Formose, à 1500 m d'altitude environ. Au contraire de Gennaeus swinhoei (son compatriote, qui se reproduit très abondamment en captivité el vit dans les jardins de Tokyo à l’état semi-sauvage), C. mikado s’est révélé un mauvais repro- ducteur en captivité, notamment les femelles, lesquelles après avoir pondu pendant leur deuxième année de vie cessent toute activité reproductrice, ce qui n’est d’ailleurs pas un cas isolé. Quelles qu’en soient les causes, C. mikado est très rare dans les collections de Faisans vivants. Pour obvier à cet inconvé- nient, j'accouplai dès 1946 un mikado © avec un ellioti G' et j'obtins des hybrides des deux sexes, pour lesquels les caractères mikado prévalaient sur les caractères ellioti, avec une telle intensité qu'on ne pouvait même pas imaginer que ces oiseaux avaient du sang d’ellioti. D'autres éleveurs ont obtenu des résultats analogues en faisant un croisement inverse, c’est-à-dire un mikado Gf et un ellioti @. Fait étonnant : ces hybrides ressemblent, plus que tout autre espèce pure ou hybride, à humiae, c’est-à-dire à l'espèce qui vit dans la région la plus éloignée de celle où vit mikado. Comme je lai déjà dit, burmanicus, voisin géo- graphique de humiae, diffère de ellioli par la couleur de la têle et du cou, qui est plus ou moins celle de humiae. C. burmanicus pourrait être considéré comme un hybride naturel humiae X ellioti, s’il n’existait une distance géogra- phique énorme entre l'aire de distribution de ces deux espèces, Source : MNHN. Paris 68 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Le résultat du croisement humiae X ellioti est lui aussi étonnant. Chez ces hybrides les côtés du cou du mâle sont blancs, comme chez ellioti ; ce caractère est le plus remar- quable, quoique les hybrides en question ressemblent beau- coup aux ellioti. Après un recroisement entre cet hybride et un mâle humiae pur, nous n'avons remarqué aucune diffé- rence saillante avec la race humiae pure. Les femelles aussi ne diffèrent pas des femelles humiae pures, tandis que les femelles hybrides du premier croisement présentent de petites taches noires à la base du cou vers la poitrine. Si on compare C. mikado avec les autres espèces, jai noté les différences suivantes : des œufs considérablement plus grands et une période d’incubation plus longue de 24 heures ; la nomenclature qui considérait mikado comme un sous-genre Cyanophasis n’est pas justifiable. Le genre Calophasis, selon nos connaissances biogéogra- phiques, comprend les espèces précitées, bien définies géogra- phiquement et non confluentes, sauf peut-être C. humiae et €. burmanicus. VALIDITÉ DU GENRE Calophasis DeLacour et BsEeBe dans leurs ouvrages respectifs et PETERS dans sa liste des oiseaux du monde, ne reconnaissent pas le genre Calophasis. En revanche celui-ci est accepté, entre autres, par OGrLVIE-GRANT et OATES qui groupent sous ce vocable en plus des Faisans à queue en échiquier (Calo- phasis proprement dit), le Faisan vénéré (Syrmaticus reevesi) et aussi le groupe des Faisans cuivrés japonais (Graphopha- sianus). Si nous nous bornons à prendre en considération la mor- phologie de ces oiseaux, nous pouvons affirmer que la queue particulièrement longue du Faisan vénéré (Syrmaticus ree- vesi) et celle des Faisans cuivrés (Copper Pheasants) pré- sentent des caractères différents. Outre que Syrmaticus reevesi (ç et Q) n’a pas les caron- cules des joues érectiles, ce qui le rend différent de toutes les autres espèces, sa queue extraordinairement longue chez le mâle l’est aussi chez les femelles. Enfin, comparées aux femelles des autres espèces, les rectrices sont pointues et non pas arrondies. Source : MNHN. Paris FAISAN DE HUME ET GENRE CALOPHASIS 69 Les Faisans du genre Graphophasianus ont un nombre de rectrices qui oscille entre 18 et 20, tandis que les S. reevesi et les autres du groupe ellioti en ont seulement 16. Or le nombre des rectrices a toujours été considéré comme un bon caractère pour la distinction des genres. Mais il y a un fait qui met fin à toute discussion, c’est la « monogonarréie » que j'ai démontrée expérimentalement entre Syrmalicus et Graphophasianus. Des expériences que j'ai failes, il résulte que les femelles hybrides, obtenues en partant de ces deux espèces, ne sont pas fécondes (tandis que les mâles le sont). Enfin elles sont morphologiquement intersexuées car leurs queues ont une longueur intermédiaire entre celles des femelles et celles des mâles reevesi. Graphophasianus est donc un genre valable du point de vue physiologique et systéma- tique, donc il doit être séparé du genre Syrmalicus. Mes expériences d’hybridation entre Syrmaticus et Gra- phophasianus ont donné de faibles résultats. De deux couples différents Graphophasianus et C. ellioti, je n’ai obtenu que deux mâles seulement, qui ont eu une vie très brève, même si l’un d’eux avait atteint l’âge adulte et avait changé les caractères intermédiaires entre les mâles des deux espèces. Le croupion de cet exemplaire montrait beaucoup de blane ce qui est un signe évident de l’hérédité paternelle, car le mâle de Graphophasianus était un soemmeringi. De toute façon la difficulté que l’on éprouve à obtenir des croisements, la faible fécondité et la faible vitalité des exem- plaires, m'ont fait conclure que l’affinité entre Graphophasia- nus et Calophasis est très médiocre. Je n’ai pas fait d'expériences d’hybridation entre Syrma- ticus et Calophasis, de toute façon mon expérience acquise après beaucoup d’hybridations me suggère que même les hybrides entre un Faisan vénéré (Syrmaticus) et un Faisan d’Elliot (Calophasis) doivent être monogonarréiques c’est-à- dire que seuls les mâles sont féconds, les femelles étant sté- riles et masculinisées. J'ai fait un croisement entre G' reevesi et Q@ ellioti, et un autre encore entre ç{ ellioti et Q reevesi dont j'attends les résultats. L'hybridisme donne des résultats étranges el imprévus, mais qui suivent de toute façon les règles de l'hérédité. Si par exemple certains caractères d’une espèce sont dominants par Source : MNHN. Paris 70 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE rapport aux caractères correspondants de l’autre espèce, du croisement d'un Polypléctron chinquis (P. bicalcaratum) avec une femelle de Faisan doré (Chrysolophus pictus) peut naître, comme j'ai pu le constater il y a quelques années, un Faisan qui juste après l’éclosion présente des caractères douteux propres au Polyplectron chinquis, mais qui par la suite devient un Faisan doré parfait. REFERENCES Bees, W. —— Pheasants, their lives and homes, Vol. 1, 2. Doubleday, New-York, 1931. Drracoun, J.— The Pheasants of the World. Country Life Limited, Lon- don, 1951. Gmcr, À. — Eterosi e mancanza di uniformità nella prima generazione ibrida fra Graphophasianus soemmeringi ijimae Dressey Syrma- ticus reevesi Gray. R. Acc. Scienze di Bologna, 1934. Oarus, E. W. — The game birds of India. Cambridge & C, Bombay, 1893. Perers, J. L. — Check-List of Birds of the World, Vol. Il. Harvard Un. Press, Cambridge, 1934. Source : MNHN. Paris DE L'EFFET DESASTREUX DE L’ENROBAGE DES SEMENCES SUR L’AVIFAUNE SUEDOISE par Sven HôRSTADIUS Institut de Zoologie d'Uppsala Secrétaire Général du X° Congrès International d'Ornithologie (Uppsala 1950) L'emploi de produits chimiques en agriculture et sylvicul- ture soulève un des problèmes les plus graves que les Protec- teurs d’Oiseaux aient jamais eu à confronter jusqu’à ce jour. Il y a deux points à distinguer : l'usage des insecticides et des herbicides d’une part, celui des semences enrobées d’autre part. Au cours de la dernière décennie, l’effet désastreux de l’enrobage des semences est devenu de plus en plus évident et c’est le moins qu’on puisse en dire ! En fait, dans plusieurs régions les espèces diminuent à une vitesse alarmante. Quoique les engrais chimiques aient été employés en Suède depuis des générations, c’est tout récemment que ces effets désastreux commencèrent à se manifester. Ceci est proba- blement dû à l'introduction de pesticides particulièrement virulents tels que aldrin, dieldrin, heptachlor, DDT, BHC et bien d’autres; mais il est dû aussi à l’usage excessif du mercure dans l’enrobage des semences. On signale qu'aux Iles Britanniques les pesticides organochlorés sont particu- lièrement nocifs. D'un autre côté il est reconnu qu’en Suède le mercure est responsable de catastrophes. L’enrobage au mercure y est d’un usage courant et récem- ment il a élé démontré que le dosage employé, quoique recommandé par l’Institut de Protection des Plantes agricoles, est beaucoup plus fort que nécessaire. En automne 1964 les rapports de cet Institut sur les effets des différents dosages révélèrent qu’un dosage à 50 % de celui communément employé obtenait les mêmes résultats. Il est déplorable que pendant des années des concentrations plus fortes que néces- saire aient été ainsi employées et il est probable que les L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris 72 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE dosages pourraient être encore réduits sans perdre de leur effet. Signalons que bien des fermiers obtiennent de bonnes récoltes sans utiliser cette technique. Au cours des dernières années de nombreux oiseaux ont été trouvés empoisonnés d’une façon ou d’une autre : morts ou paralysés. Comme dans quelques cas la présence du mer- cure fut décelée à la suite de l'analyse par l’Institut Vétéri- naire de Stockholm, et comme d’autre part certaines espèces diminuaient en nombre d’une façon alarmante, la Société Ornithologique de Suède commença une enquête dès jan- vier 1964, notamment auprès de ses membres, afin de con- naître les changements observés quant aux taux des popula- tions et d'obtenir des informations sur les cas sûrement imputables aux pesticides. Les résultats furent publiés par G. Orrercind et I. LENNERSTEDT, dans la revue de cette société, Vär Fagelvärld, vol. 23, 1964, pp. 363-415, avec un résumé en anglais. Les informations concernant les insecticides et les herbi- cides sont aussi alarmantes, mais les principaux dégâts paraissent êlre dus à l’enrobage des semences à base de mer- eure. Dans sa première manifestation l’empoisonnement rend l'oiseau beaucoup plus familier au point qu’il peut être plus facilement attrapé, ensuite les pattes sont paralysées et enfin l'oiseau meurt dans des convulsions. On ne doit jamais perdre de vue qu’il peut y avoir, comme cause d’un déclin sérieux d’une espèce, d’autres facteurs que les pesticides. Une culture plus rationnelle peut considérable- ment changer l'accès aux lieux de nidification et d’alimenta- tion pour beaucoup d'oiseaux. Le nombre décroissant des chevaux et des têtes de bétail peut également avoir son impor- tance. En Suède, le déclin de l’Alouette lulu (Lullula arborea) peut être imputé à la mise sous forêt des petits champs entou- rés de bois, où elle préfère nicher. Mais, il ne peut y avoir aucun doute : l'énorme déclin numérique de quelques espèces est en relation avec l'emploi des pesticides dans les grandes régions agricoles, car dans les régions d'agriculture moins intensive comme dans les pays boisés ou dans les archipels les populations ont été moins affectées. Le Bruant jaune (Emberiza citrinella), le Bruant ortolan (Emberiza hortulana) et le Faucon crécerelle (Falco tinnunculus) ont pratiquement disparu des grandes régions de culture. On signale la dispari- tion graduelle du Bruant jaune dans presque toutes les Source : MNHN. Paris EFFET DÉSASTREUX DE L'ENROBAGE DES SEMENCES 73 régions de Suède. Un cas de patte paralysée a été observé et du mercure a été trouvé chez un spécimen. De semblables constatations ont été faites pour le Bruant ortolan ; dans un cas les oiseaux, sûrement paralysés par le poison, furent observés dans des champs où l’on avait utilisé des semences enrobées. La Crécerelle était communément aperçue survolant les champs des régions cultivées ; mais maintenant elle a tota- lement disparu. Pourtant elle n’a pas diminué dans les archi- pels. Des cas d’empoisonnement par mercure ont en revanche été confirmés. Le Corbeau freux (Corvus frugilegus) a subi une diminu- tion considérable. De nombreuses colonies ont disparu des régions agricoles et cette diminution est estimée environ à 75 %. 11 n’est pas sûr que le Bruant proyer (Emberiza calan- dra) continue à se reproduire en Suède. De même le Moineau domestique (Passer domesticus) et le Moineau friquet (Pas- ser montanus) ont considérablement diminué en beaucoup d’endroits. Ces deux oiseaux, facilement observés dans les cours des fermes, ont fait l’objet de nombreux rapports qui constatent la paralysie chez des individus nourris de semences enrobées, et signalent de multiples jeunes trouvés morts au nid. D'autres rapports font état de Pigeons ramiers (Columba palumbus) paralysés où morts, et ceci souvent en relation avec l’ingestion de semences enrobées. Des traces de mer- cure ont été trouvées au moins 3 fois dans les oiseaux morts. Presque tous les rapports sur la Perdrix grise (Perdix per- dix) soulignent une importante régression de la population. Peut-être même peut-on dire que la population encore exis- tante en Suède méridionale n’atteint guère plus de 10 à 50 % de ce qu’elle était il y a 10 ans. Souvent le poussin meurt dès son jeune âge. Pourtant en ce qui concerne la Perdrix, l’en- robage ne peut être accusé d’être le seul facteur dangereux. Il est évident que les herbicides et pesticides jouent aussi un rôle important mais l’analyse a trahi la présence de mercure dans plusieurs cas. Il en est de même pour nombre de Fai- sans trouvés morts, mais la régression de l'espèce n’est pas aussi certaine que pour la Perdrix. Dans quelques endroits de nombreux Faisans ont été trouvés morts où manifestant des signes typiques d’empoisonnement. Dans une région où le printemps fut particulièrement sec, le blé enrobé demeura tel plusieurs semaines avant de pousser ; un déclin des Fai- Source : MNHN. Paris 74 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE sans se fit alors sentir l’hiver suivant, en même temps que le nombre d'individus tués par les oiseaux de proie était par- ticulièrement élevé. Les oiseaux de proie, en tant que dernier échelon du cycle trophique, sont spécialement intéressants. Nous avons déjà signalé la disparition désastreuse de la Crécerelle ; le Faucon pélerin (Falco peregrinus) a lui aussi considérablement dimi- nué, mais aucun cas d'empoisonnement n’a pu être vérifié. Les informations touchant la très petite population de Pygargue à queue blanche (Haliaëtus albicilla) sont particu- culièrement alarmantes. L'un d’eux, vu les jambes paralysées, mourut le lendemain ; un autre trouvé mort révéla des traces de mercure dans le foie et les reins. Mais, ce qui est encore plus grave, plusieurs rapports ont constaté l’insuccès des cou- vées. Presque aucun jeune n’a pu être élevé, ils mouraient même avant que de pouvoir voler ; et dans beaucoup de cas, l'instinct de reproduction semble avoir été inhibé. Enfin un œuf d’un nid abandonné fut analysé et contenait 3,5 mg/kg de mercure. De semblables constatations concernant des régressions de population, une réduction de la fréquence de reproduction et quelques cas dûment reconnus d’empoisonnement par le mer- cure ont été faites pour l’Autour des palombes (Accipiter gen- tilis), l’Epervier d'Europe (Accipiter nisus), et la Buse variable (Buteo buteo). Enfin on signale encore des taux de reproduction parti- culièrement bas ainsi qu'une forte mortalité juvénile chez le Hibou grand-duc (Bubo bubo), le Hibou moyen-duc (Asio otus) et la Chouette hulotte (Strix aluco). En ce qui concerne cette dernière, deux jeunes furent trouvés morts 10 jours après l’ensemencement des champs environnants, ils ne conte- naient pas moins de 270 mg/kg de mercure dans le foie et les reins. Les faits ci-dessus ne donnent qu’une idée du matériel d'étude présenté par OrrerLiND et LENNERSTEDT, mais mon but étail de souligner l'immense menace qui résulle pour beaucoup d'espèces d'oiseaux de l'emploi généralisé du mer- cure dans l’enrobage des semences en Suède. Mais ce n’est pas tout, le poison s’accumule dans le cycle trophique au niveau où interviennent les prédateurs et l’homme. Tout récemment un rapport préliminaire dont il fut fait état dans les jour- naux signale que l’Institut Vétérinaire de Suède (qui fait les Source : MNHN. Paris EFFET DÉSASTREUX DE L'ENROBAGE DES SEMENCES 75 analyses) a trouvé chez les Faisans tués au cours de la sai- son de chasse jusqu’à 800 et 900 fois plus de mercure qu'il n'en était toléré dans la nourriture par l'Organisation Inter- nationale de la Santé ! Nous avons donc atteint la limite au- delà de laquelle l'usage excessif des poisons va devenir un réel danger pour l’homme et cela malgré les cris d'alarme des protecteurs. Espérons que des faits aussi parlants ouvri- ront les yeux de ceux qui se refusent à voir le danger et les amèneront à une contre-réaction dont bénéficieront également les oiseaux. De nouveaux faits survenus depuis la remise du manus- crit méritent d’être mentionnés ici. Des analyses opérées par l’Institut Vétérinaire afin de déceler la quantité de mercure contenu dans le foie et les reins d'oiseaux trouvés morts ou mourants, 50 % ont révélé s'inscrire entre les chiffres suivants : Faisans et Perdrix : 28-150 mg/kg; Pigeons : 8-45; Corvidés : 29-110; Frin- gilles : 11-136 ; Rapaces diurnes : 6-100 ; Rapaces nocturnes : 4-270 mg/kg. Six œufs de Pygargue (Haliaetus albicilla), pro- venant de régions différentes, contenaient de 3,5 à 11 mg/kg. Du mercure a élé également trouvé chez le renard, la martre, le putois, mais non chez le chevreuil et le lièvre. Même pour ces oiseaux tués à la chasse ou par accident et ne montrant aucun signe de maladie, les analyses se sont montrées posi- tives. Sur 39 oiseaux se nourrissant de graines, on a trouvé du mercure dans 26 d’entre eux, et 50 % de ceux-ci présen- taient de 1 à 39 mg/kg dans le foie. Sur 70 Autours et Buses, 67 ont donné des analyses positives ; 50 % des Autours don- naient de 6 à 53 mg/kg et les Buses de 2,1 à 65 mg/kg dans le foie. Des expériences faites sur les Faisans nourris de graines traitées au mercure ont prouvé une diminution du nombre d'œufs couvés avec succès. L'Institut Vétérinaire a prouvé statistiquement la valeur de ces analyses et expé- riences. Il est donc probable que le grand responsable de ce drame est l'enrobement des graines au mercure. Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS OF FOREST AND SAVANNA by David Lack and R. E. MorEAU Edward Grey Institute, Oxford THEORIES OF CLUTCH-SIZE As is well known, the passerine birds of the tropics tend to have much smaller clutches than those of the temperate zones (Moreau 1944). On the theory supported by Lack (1947), this is because, in general, tropical birds can collect less food for their young than those of temperate regions, and the usual reason suggested for this is that in summer the diurnal birds of north-temperate regions have about 1 ?/, times as many hours each day in which to collect food as tropical species. But this difference, though undoubtedly important, is inadequate in itself Lo account for the whole difference, since in various passerine families the temperate- zone species lay clutches of twice the size of their tropical congeners. Asuwoze (1961), accepting the idea that tropical species not only do not, but cannot, raise so many young as north- temperate birds, postulated an additional reason for their inability. Among north-temperate birds, those which are sedentary are greatly reduced in number in winter by food shortage, while those which migrate are presumably reduced by the dangers incurred on migration. As a result, the bree- ding populations of such species tend to be low in relation to the seasonal flush of insects and other foods for their young in spring and summer. In relatively uniform tropical habitats on the other hand, there may be no marked seasonal food shortage, and in any species that is limited by its food supplies, the adult numbers will probably be close to the limit set by food throughout the year, so that there will not at any time be a big surplus of food out of which young L'Oisean et R.F.O., V. 35, 1965, n° spécial. Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 77 birds can be fed. Hence, the number of young that can be raised will be small. AsamoLe’s postulate depends on the truth of two general theories, first that the numbers of passerine birds are limited in a density-dependent way by their food supplies outside the breeding season, and secondly that the clutch-size of each species has been evolved to correspond with that brood-size from which the food supply permits the greatest number of surviving young to be raised (Lac 1954). Both theories have, of course, been questioned, but there is not space here to discuss them in general. It is relevant, however, to note that both Sxurcx (1949) and WaGner (1957, 1960) have argued that, while the above interpretation of clutch-size might perhaps hold in temperate regions, it is not applicable to tro- pical passerine species. SkuTcH in particular claimed that tropical passerine species could normally collect much more food than is needed to raise broods of only two young. He therefore suggested that natural selection might operate against the evolution of a larger brood because the latter would entail more feeding visits by the parents, thus increa- sing the risk of the nest being found by predators. ASHMOLE’s postulale also depends on a third point, that the amount of food which can be brought by a pair of birds to their young is less when the density of breeding pairs is higher. This seems reasonable on general grounds. Also the Great Tit Parus major lays a smaller clutch at higher popu- lation densities (KLurver 1951, Lack 1958), presumably because it cannot raise so many young at higher densities, and at least in one year near Oxford there was a much lower production of young in one wood with an exceptionally high density of breeding pairs than in another wood with a nor- mal population density (PERRINS 1963). It occurred to us that ASHMOLE’s view could be tested by a comparison of cluich-size within the tropics, between species breeding in evergreen forest, which provides the most uniform conditions, and those breeding in savanna and other drier habitats with a restricted rainy season. In the latter habitats, during the long annual drought, the trees to some extent lose their leaves while the ground cover becomes desic- cated and is often removed by fire. Hence, though measure- ments have not been made, it seems certain that the food supplies diminish, and consequently that the birds become Source : MNHN. Paris 78 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE much reduced in numbers between successive breeding sea- sons (cf. Morez and BouruièrE 1962). Since both forest and savanna are in the tropies, the complication of a difference in daylength is avoided. Actually Moreau (1944) had already found that average cluiches tend to be larger in savanna than forest, but much more extensive data are n0W available for the clutches of African species, suggesting that a fresh enquiry would be rewarding. THE AFRICAN DATA For the purposes of this comparison, We decided to limit ourselves to passerine species, and among these especially concerned ourselves with those families in which cluteh-size records were available for at least five species in forest and five outside forest, in order to avoid possible bias due to one or two exceptional species. The main sources of information are Moreau (1944), who had already included all published records up to that date for East Africa, and CHapin (1953, 1954) primarily for the Congo but also summarising infor- mation from elsewhere. One family, the babblers Timaliidae, was inadequately represented in both these sources, so fur- ther information was taken from PrAED and GRANT (1955), while to obtain records from arid country, we consulted Ancner and Gopwan (1961) for Somaliland. For the Ploceidae only, we undertook a wider search, covering all the standard regional ornithologies of tropical Africa. Data for Africa were restricted to the tropical mainland, records from islands off the coast and also from South Africa being omitted. This was because islands do not provide eco- logical conditions comparable with the mainland, while clutches are in general higher in South than tropical Africa (Moreau 1944). The full list of species utilised and their respective clutch-sizes are set out in Appendix 1, while the method of assessing the average clutch-size for each family is set out in Appendix 2. The numerical assessment of clutch- size from the published data was a difficult and somewhat arbitrary procedure, but quantitative tabulation was essential for our purpose, and as we have published our figures, other workers can see what we have done. For a number of species later knowledge may show that our figures are somewhat wrong, while for others there are as yet too few records for Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 179 a firm conclusion, but there is no reason to think that any errors are in one direction rather than another, so that they should not affect the general trends discussed here. The main findings for tropical Africa are summarised in Table 1, of which the upper section fully confirms MOREAU’S earlier conclusion that, in general, clutch-sizes are lower in evergreen forest than in habitats outside it, most of the latter having deciduous vegetation and a well-marked dry season. Of the eight families for which the averages rest on at least 5 species in each habitat, the non-forest species have an ave- rage clutch-size of rather over half an egg more than the forest species in the Estrildidae, Lanïidae, Ploceidae, Tima- lidae and Turdidae, and there is a smaller difference in the same direction in the Sylviidae, but virtually no difference in the Muscicapidae or Nectariniidae. The figures for two families need qualifying. The higher average cluteh-size of Sylviidae outside than in forest is due almost entirely to the 23 species of Cisticola, for which the average clutch-size was 2.9, and if this genus were omitted, on the grounds that none breed in forest, the average for the non-forest species of Sylviidae would be reduced to 2.4, only a little larger than for the forest species. There are, of course, various other genera of Sylviidae which are restricted to one habitat or the other, and there are no grounds for excluding Cisticola rather than any of these others except that it contains so many species. Similarly in the Ploceidae, all 15 species of Bubalornithinae and Passerinae for which the clutch-size is known breed outside forest and have an average clutch-size of 3.5, whereas 44 species of Ploceinae which breed outside forest have an average clutch-size of 2.5, as compared with an average of 2.1 for the 8 forest species of Ploceidae (all in the subfamily Ploceinae). Thus, even when the comparison is restricted to the Ploceinae, the forest species have a smaller average clutch than those outside forest. Suggestive evidence of a trend in the same direction was found in those families for which fewer than five species were available for comparison in both habitats. In the Stur- nidae (starlings), three forest species have an average clutch- size of 2.7, compared with an average of 3.5 for 17 species outside forest. Again, in the Hirundinidae (swallows), Dicru- ridae (drongos), and Fringillidae (cardueline finches), the 6 Source : MNHN. Paris 80 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE single forest species for which data are available has a smaller clutch than the average for respectively 17, 1 and 16 species breeding outside forest. But the 21 species of Pycnonotidae (bulbuls) which breed in forest lay clutches of two, like the single species outside forest. It is also suggestive that, except for the estrildines and carduelines, almost all passerine species breeding in forest have clutches of wo, whereas the average clutch-size is three eggs or more in nearly all the other tropical African passerine families which breed entirely outside forest, including the Alaudidae (larks), Corvidae (crows), Paridae (tits), Certhidae (creepers) and Motacillidae (pipits, wagtails), while it exceeds two, though under three, in the Oriolidae (orioles). In view of these general trends, it is surprising to find that nearly all the tropical African species of Muscicapidae have clutches of two, irrespective of habitat, the few excep- tions being noted in Appendix 1. Again, none of 39 species of Nectarinidae has been recorded as laying a clutch larger than two eggs, but in this family a few species lay only one egg. Nearly all such records are from the eight species for- merly kept together in the genus Nectarinia, sensu striclo, five of which, though not the others, are montane. Most other African sunbirds have since been brought within this genus (which was the earliest to be described in the family), but for convenience we have retained the former classification in Appendix 1. The latter also includes a few other species in which a clutch of one has been recorded. If ASHMoLE’s view is correct, then it might also follow that the species breeding in extremely arid habitats should lay larger clutches than those which, though also breeding out- side forest, frequent less extreme habitats. There are records for a sufficient number of species to make such a compari- son possible for two families, the Alaudidae and Sturnidae, and the lower section of Table 1 shows that, in both, the ave- rage clutch is over 1 ‘/, eggs lager in the very arid habitat of Somaliland than elsewhere in tropical Africa outside forest. AN AMERICAN COMPARISON SkurcH (1954, 1960), working mainly in humid forest and cultivated land in Costa Rica, and MarCHANT (1960), working in the extremely arid Santa Elena peninsula in Ecuador, Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 81 obtained sufficient clutch-records to permit us to make simi- lar comparisons in relation to habitat for two tropical Ameri- can families. As shown in Table 2, in both the Tyrannidae and the Fringillidae, the average clutch-size was rather larger for the species breeding in forest clearings and other more open habitats in Costa Rica than for the local forest species, and larger again for those breeding in the Santa Elena penin- sula than in open habitats in Costa Rica. Hence both diffe- rences are in the same direction as in the equivalent habitats in Africa though each average is as yet based on rather few species in America. DISCUSSION As mentioned in the introduction, ASHMOLE postulated that species living in a habitat with little seasonal change during the year would probably be close to the limit of num- bers set by food almost throughout the year, and so would find it hard at any time to find enough additional food to raise young and hence would evolve small clutches. We pre- dicted from this theory that the average clutch-size should be lower in tropical forest than in savanna, and this was found to be true in tropical Africa, while there was some confirmation from tropical America also. It does not necessa- rily follow from this that ASHMOLE’s postulate is right, but the fact that it holds for a variety of passerine families in two continents is highly suggestive, and it may be added that ASHMOLE was evidently unaware of this difference between forest and savanna species when he wrote, Even so, there may well be additional factors modifying the size of brood which different species can raise, and in his context it is almost certainly important that the forest species of Ploceinae, and some of the forest species of Estrildidae, are insectivorous, whereas those of savanna eat mainly seeds. Doubtless fur- ther knowledge would show that there are other differences in ecology which help to account for the differences in clutch- size between forest and non-forest species in particular fami- lies. But as the generalisation holds for nearly all the fami- lies investigated here, it is reasonably certain that a general factor is involved in addition to any particular factors affec- ting some but not other species. It should be added, however, that while in nearly all the Source : MNHN. Paris 82 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE families investigated, the species breeding outside forest have larger clutches than those in the forest, the average diffe- rence is only about half an egg, i.e. about one quarter as much again. This is a much smaller difference than might, perhaps, have been expected, at least from the impression that the observer gets of the relatively uniform conditions in forest on the one hand and the temporary flush of foods in savanna on the other hand. One would have supposed that the seasonal difference in food resources between the two environments would be much greater than in the proportion of 4:5, but actual measurements have not been made, and this superficial impression of the ecological situation might be fallacious. Alternatively there may be other factors modi- fying clutch and brood-size. Since this paper provides support for ASHMOLE’S view, it also provides support for the view that tropical species, like those of north temperate regions, raise as large broods as they are capable of feeding, and that they have evolved clutches to correspond. But this evidence is as yet purely circumstantial, and the experimental test of providing a tro- pical passerine species with a larger brood than its normal elutch-size has not been undertaken. SUMMARY 1. ASHMOLE suggested that one reason why tropical spe- cies have smaller clutches than those of temperate regions is that the tropical species live under nearly uniform conditions throughout the year, so that their numbers remain suffi- ciently close to the limit set by food for them to find it hard at any time Lo feed a brood, and accordingly, they have evol- ved small clutches. Those of temperate regions, on the other hand, being reduced heavily in numbers in winter and there- fore scarce relative to their food supplies in spring, can raise larger families and evolve larger clutches. 2. This difference should apply equally within the tro- pics to species breeding in evergreen forest and savanna res- pectively, as the latter experience a period of food shortage near the end of the dry season. In fact such a difference holds in 6 out of 8 African passerine families in which the clutch- size is known for at least 5 species in each habitat, and there Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 83 is also suggestive evidence for it in four other African fami- lies represented by fewer species in one or other habitat, and in two tropical American families. 3 No difference in clutch-size between forest and savanna species was found in Muscicapidae or Nectarinidae. 4. In each of four families, two in Africa and two in America, the average clutch-size was larger in an extremely arid habitat (han in less extreme conditions outside forest. TABLE 1 Average clutch-size in different habitats in tropical Africa (number of species in brackets) TROPICAL AFRICA Family outside forest forest Estrildidae (waxbills) 3.7 (10) 4.3 (32) Lanïidae (shrikes) 2.0 (5) 2.7 (26) Muscicapidae (lycatchers) 2.0 (15) 2.2 (22) Nectariniidae (sunbirds) 1.9 (11) 1.7 (83) Ploceidae (weavers) 2.1 (8) 2.8 (59) Sylviidae (warblers) 2.2 (14) 2.6 (51) Timaltidae (babblers) 2.0 (5) 2.7 (0) Turdidae (thrushes) 2.0 41) 2.7 (26) other arid non-forest Somaliland Alaudidae (larks) 21 (6) 3.7 (14) Sturnidae (starlings) 3,0 (12) 4207) Notes : Among the non-forest Ploceidae, 44 species of Ploceïnae had an average clutch-size of 2.5 and 15 species of Bubalornithinæe and Passerinae of 3.5. Among the non-forest Sylviidae, 23 species of Cis- ticola had an average clutch-size of 2.9 and 28 other species of 24. TABLE 2 Average clutch-size in different habitats in tropical America (number of species in brackets) Family Mainly humid Costa Rica Very arid forest others S.W. Ecuador Fringillidae (finches) 1.9 (3) 2.2(5) 3.0 (6) Tyrannidae (tyrant-flycatchers) 2.1 (6) 2.5 (20) 2.9 (8) Source : MNHN. Paris 84 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE APPENDIX CLuTrGH-SIZE RECORDS USED FOR TABLES 1 AND ? Note : À single number or two numbers joined with a hyphen refer to general statements on clutch-size, e.g. 3 means normal elutch 3, 4-5 means normal clutch 4 or 5. In such cases very unusual clutch- sizes are omitted. À number with another in brackets means that the latter is less frequent, e.g. 34), while if the second number is in double brackets it is unusual. One number followed by a stroke and then a second number, e.g. 1/4, means one record of a cluteh of 4; this notation is used only when very few records (which might perhaps be atypical) have been published. Within each family the genera, and within each genus the species, have been listed alphabetically. For con- venience we have followed the nomenclature used by the authors cited, basically CnariN (1953, 1954) and Moreau (1944). Two specific names joined by + mean that two forms formerly classified as separate spe- cies are now united in one. TROPICAL AFRICA Estrildidae Forgsr : Inseenvorous Nigrita bicolor, 1/4, 1/5, canicapilla 4, fusconota 1/6, Parmoptila mwoodhousei 3-4, Pholidornis rushiae 2; Orners : Cryptospiza reichenowi + australis (2) 3, Salbadori 2/4, Estrilda atricapilla 4-5, Hypargos niveoguttatus 1/3, Spermophaga rufi- capilla 1/3. Now-Foresr : Amandava subflava 4-5, Clyptospiza monteiri 6, Coc- copygia quartina + melanotis (3)4, Estrilda astrild 4-5, melpoda 5- nonnula 4-6, paludicola 4-5, troglodytes 6, Euodice cantans 1/5, 1/6, Granatina ianthinogaster 3-5, Lagonosticta jamesoni 3-4, larvata + togoensis 3-4, perreini 3-4, rara 1/4, rubricata 3-5, rufopicta 3/4, sene- gala 3-5, troglodytes 6, Lonchura bicolor + nigriceps 4-6, cucullata 4- fringilloïdes 1/6, Nesocharis ansorgei 1/2, Ortygospiza atricollis 6, locustella (3)5, Pirenestes ostrinus (3)4, Pytilia afra 3-4, melba 4-5, phoenicoptera 3/4, Spermophaga haematina 3, Uraeginthus angolen- sis 3-6, bengalensis 4, cyanocephalus 1/3. Lanïdae Forest : Chlorophoneus bocagei 1/2, nigrifrons 1/2, Laniarius leu- corhynehus 2, Malaconotus lagdeni 1/2, Nicator chloris 2. Now-Foresr : Antichromus minutus 2(3), Corvinella corvina 4(5), Chlorophoneus sulfureopectus 2, Dryoscopus cubla 2-3, gambensis 2, prin- glii 1/9, Eurocephalus anguitimens 2-4, Laniarius ergthrogaster 9, fer- rugineus 2(3), funebris 3, lühderi 2, ruficeps 1/3, Lanius cabanisi 3-4, collaris 2-4, excubitorius 3-4, mackinnoni 2(3), somalicus 4, souzae (2) 3, Malaconotus cruentus 2-3, poliocephalus (2) 3, Nilaus afer 2, Prionops cristata + poliocephala 4-5, retzii 2, Tehagra australis 2 (3), jamesi 2-3, senegala 2 (3). Muscicapidae Fons : All 2 : Alseonax epulatus, seth-smithi, Dioptrornis fischeri, Dyaphorophyia blissetti, castanea, chalybea, Fraseria ocreata, Pedi- Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 85 Lorhynchus comitatus, Terpsiphone rufocinera + nigriceps, Trochocercus albiventris, albonotatus, cyanomelas, nigromitratus, nitens ; also Alseo- naz adustus + minimus (1) 2(3). Nox-Fongsr laying 2 : Alseonat aquaticus, cassini, Artomyias fuli- ginosa, Batis minor, molitor, Bias musicus, Bradornis griseus, micro- rhynchus, Elminia albicauda, longicauda, Hyliota flavigaster, Hypodes cinerea, Melaenornis edolioides, Muscicapa gambagae, Parisoma lugens, Platysteria cyanea. Now-Fonxsr others : Batis orientalis 1/2, 1/3, Bradornis pallidus 2-3, Hyliota australis 1/3, Melaenornis pammelaina 23, Myopornis bôhmi 1/4, Terpsiphone viridis + perspicillata 9-3. Nectarinidae Fonrsr (a) laying 2 : Anthreptes aurantium, tephrolaema, Cinnyris minullus, superbus, Cyanomitra batesi, cyanolaema, olivacea, seimundi ; (b) others : Anthreptes gabonicus ((1))%, yokanae 1/1. Nox-Fonesr laying 2 : Anthreptes anchietae, collaris, longuemari, Chalcomitra amethystina, angolensis, fuliginosa, rubescens, senegalensis, Cinnyris chloropygins, coccinigaster, cupreus, johannae, osea, reiche- nowi, shelleyi, venustus, Cyanomitra verticalis, Hedydipna platura (ARCHER and Gopmax quoted 3 for the Sudan without citation, so we have ignored it). Nox-Fonesr others : Chalcomitra hunteri 1/1, senegalensis ((1))2, Cinnyris albiventris 2/1, afer 2/1 (2 sometimes 1 even in South Africa), bifasciatus (1) 2, habessinicus 1 ((2)), Cyanomitra reichenbachi 2/1, Nec- farinia erythrocerca 1(2), famosa (1)2, johnstoni 1, kilimensis 1(2), nectarinioides 1-2, pulchella 1-2, reichenowi 1, focazzae 1. Ploceidae Forgsr PLOGENAE : Malimbus malimbicus 2, nitens 2(3), scutatus 2, Ploceus alienus 2, bicolor 2-3, insignis 2, melanogaster 2, nigricol- lis (D) 2. Nox-Fonesr PLOGEINAE : Amblyospiza albifrons 2-3, Euplectes afra 2-4, albonotatus 2-3, anomala 2/2, ardens 2-8(4), axillaris 2-3, capensis 2-3, gierowii (2)3((4)), hartlaubi %, hordeaceus 2-4, jacksoni 2-3, macrourus 2-4, nigroventris 2-3, oryx 2-4, progne 2-4, Malimbus rubri- ceps 3, Ploceus aurantins %, baglafecht X3), bertrandi 2, bojeri 2, cas- taneiceps 283), castanops 23), eucullatus X3), galbula 3, heuglini 2, intermedius 2, jacksoni %3), Inteolus 2-3, melanocephalus 23), niger- rimus 23), ocularis 2(3)), pelzelni 2-3, rubiginosus 3-4, spekei 3, subau- reus 2-4, superciliosus 3(4), faentopterus (3), velatus 3), vitelli- nus 2-3, œanthops 2, æanthopterus 2, Quelea quelea 3, cardinalis 2-3, ergthrops 2-3, Nox-Fonesr BusaLornirHinar : Bubalornis albirostris 2-3, Dinemellia dinemelli 3-4, Nox-Fonesr PAsseNAr : Histurgops ruficauda 3, Passer castanopte- rus 5-7, eminibey 3-4, griseus 3-4, iagoensis 3-4, luteus 3-6, Petronia brachydactyla 3-5, superciliaris 3-4, Plocepasser mahali 2-3, supercilio- sus 2-3, Sporopipes frontalis 3-4, Pseudonigrita arnaudi 3, cabanisi 2-4. Sylviidae Fonesr : Apalis binotata 2, jacksoni 1/2, pulchra 23), thoracica 2/2, Camaroptera chloronota 1/2, superciliaris 1/3, Hylia prasina 1-2, Pri- Source : MNHN. Paris 86 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE nia bairdii 3, Sathrocereus mariae 2, Seicereus ruficapilla 3, umbro- virens 1/3, Sylvietta denti 1, leucophrys 1/2, virens 263). Now-Foresr other than Cisticola : Apalis flavida 2-4, rufifrons 1/3, 1/4, Acrocephalus baeticatus 2, Bradypterus brachypterus 2, cinnamo- meus 1/2, 1/3, Calamocichla gracilirostris A3), parva 2, rufescens 2-3, Camaroptera brevicaudata %3), simplex 1/3, 1/4, Chloropeta natalen- sis 24), similis 1/2, Eminia lepida 23), Eremomela canescens 1/1, icte- ropygialis 1/3, griseoflava 2/2, Heliolais ergthroptera 1/2, Melocichla mentalis 2, Phyllolais pulchella 2, Prinia leucopogon 2-3, mistacea 3(4), somalica 4, subflava 2-4, Schoenicla brevirostris 2, Syloietta bra- chyura 2, isabellina X3), ruficapilla 1/2, whytit 2. Nox-Foresr Cisticola : angusticauda 3-4, anongma 2, aridula 1/4, ayresii 3-4, brachyptera (2)3, brunnescens 2-3, bulliens 3, cantans 3-4, chiniana 3, chubbi (2)3, dambo 23), emini 3, ergthrops 2-3, eximia 2- galactotes 3-4, juncidis 3-4, lateralis 2-3, natalensis 2-3, pipiens 3- robusta 2-3, rufilata 2-3, linniens 3, woosnami 2. Timaliidae Forest all 2 : Malacocincla cleaveri, fulvescens, rufipennis, Pseu- doalcippe abyssinica, Ptyrticus turdinus. Now-Foresr : Argya aylmeri 2-3, rubiginosa 2-4, Turdoides hypo- leuca 3(4), jardinei 1/3, leucopygius 3, melanops 2-3, plebejus 2-3(4), reinwardit 2, fenebrosus 2-3. Turdidae Fonesr : Cossypha bocagei 1/2, cyanocampter 2, Geokichla piag- giae 1/2, Illadopsis rufipennis 1/2, stictigula 2, Neocossyphus rufus 1/2, Pogonocichla stellata 23), Sheppardia cyornithopsis 1/2, sharpei 2, Tur- dus abyssinieus 2, olivacens 2(3). Now-Foresr : Cercomela familiaris 3-4, melanura 3, Cercotrichas podobe 2-4, Cichladusa arguata 2, guttata 2, ruficauda 1/2, Cossypha caffra 2-3, heugliñi 2-3, niveicapilla X3), natalensis 2-3, Erythropygia barbata 2-3, galactotes 1/3, hartlaubi 1/2, leucophrys 2-3, Myrmecoci- chla arnotti 34), nigra 1/2, 1/3, Monticola angolensis 3-4, rufocine- rea 1/4, Oenanthe albifrons 2/3, phillipsi 2/3, 1/4, pileata 3, schalowi 3, Saxicola torquata (2)3-4, Thamnolea cinnamomeiventris 2, Turdus libo- nyanus (2)3, ludoviciae 2/2. Alaudidae SomattLaNn : Ammomanes deserti 3-4, Calandrella rufescens 4-5, Certhilauda alaudipes 3, hammertoni 3, somalica 1/3, 1/4, Eremopterix nigriceps 4, signata 46), Galerida cristata 4-5(6), theklae 4-5, Hetero- mirafra ruddi 3, Mirafra cantillans 4, collaris 3, gilletti 3, Pseudalae- mon freemantlit 1/3, 1/4. Oruer Troricaz : Calandrella cinerea 2, Eremopterix leucopareia verticalis 2-3, Heterocorys modesta 2/1, Mirafra africana 2-3, rufocii namomea + fischeri 3). Sturnidae Fonesr : Lamprotornis corruseus 3, purpuropterus (3), Poeoptera lugubris 1/3. Somazanp : Buphagus ergthrorhynchus 2-3, Cosmopsarus regius 4-6, Lamprocolius chalybeus 4, Onychognathus blythii 45), Speculipas- tor bicolor 3-4, Spreo albiceps 5-6, shelleyi 4-5(6), superbus 4. Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 87 Ormen Now-Fonesr : Buphagus africanus 2-3, erythrorhynchus 2-3, Cinnyricinclus leucogaster 2-3, Creatophora cinerea 2-3, Lamprocolius chalybeus 2-46), splendidus 2-3, Lamprotornis caudatus 24, Onycho- gnathus morio 3, fenuirostris 3-4, Spreo hildebrandi 3-4, puicher 3-5, superbus 2-4. TROPICAL AMERICA Fringillidae Fongsr : Arremon aurantirostris 2, Atlapetes lorquatus 2, Cyano- compsa cyanoides (1)2, Orex Wooptaxn : Saltator maximus 2. Now-Fonesr : Arremonops conirostris (3)), Saltator albicollis 2, Sporophila aurita ?, torqueola 3), Tiaris olivacea 2-3. Sanra tea : Neorhynchus peruvianus (2)3, Pheucticus chrysope- plus 2-4, Poospiza hispaniolensis 3-4, Rhodospingus cruentus 4(4), Spo- rophila telasco 2-3, Volatinea jacarina 3. Tyrannidae Forest : Myiobius sulphureïpygius %, Onychorhynchus mexicanus ?, Pipromorpha oleaginea (2)3, Platyrhinchus coronatus 2, Rhynchocyclus brevirostris 2, Terenotriccus erythrurus 2/2. Fonesr AND oPex : Megarhynchus pitangua 2-3. Nox-Fonrsr (CENTRAL America) : Capsiempsis flaveola 2, Contopus cinereus 2/3, Elaenia chiriquensis 2, flavogaster 2, Legatus leucophaius 26), Myiarchus tubereulifer 2/4, Mytobins atricaudus 1/2, Myiodynastes luteiventris 1/3, maculatus (2)3, Myiophobus fasciatus ?, Myiozetetes cayanensis 2-3, granadensis (2)3, Similis (2)3-4, Sayornis nigricans 2-3, Serpophaga cinerea 2, Todirostrum cinereum (2)3, syluia 2, Tolmomyias sulphurescens 2-3, Tyranniscus vilissimus 2, Tyrannus melancholi- cus 2-3. Sanra ELEnA : Camptostoma obsoletum X3), Elaenia leucospodia (2)3, Euscarthmns melorgphus 2, Muscigralla brevicauda 3-5, Myiodynastes bairdi 465), Phaeomyias murina ?, Pyrocephalus rubinus (2)3, Tyrannus niveigularis (2)3-4. APPENDIX 2 METHOD OF CALCULATING AVERAGE CLUTCH-SIZES FOR TABLE 1 In calculating the average clutch-size for each family in each habi- tat, the average for each species recorded in Appendix 1 was given equal value, ie. each was reckoned as one figure irrespective of the number of records on which it was based. The score used for the clutch-size of each species in caleulating the average is best illustrated by actual examples thus, 3 was scored as 3, 2-3 as 2.5, 23) as 2.25, (23 as 2.75, 2,3(4) as 2.75, 2(3)) as 2.1. The total for all the species in each group was then summed and divided by the number of species involved. Hence the average for each group is the average of the averages for all the species involved. Source : MNHN. Paris 88 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE RESUME 1°) ASHMOLE suggère qu’une des raisons pour lesquelles les pontes d'espèces tropicales sont plus petites en nombre que celles des espèces vivant en région tempérée tient au fait que les espèces tropicales vivent loute l’année dans des conditions à peu près uniformes. Le nombre d'individus reste donc voisin des limites permises par la quantité de nourriture à leur disposition. Il en résulte qu'il leur est difficile d’élever une nichée, alors que dans les régions tempérées le nombre des individus étant fortement réduit par l'hiver, donc faible pour la nourriture offerte au printemps, ces derniers peuvent élever des familles plus importantes, d'où ponte plus forte. 2°) Cette différence se retrouve entre les espèces tropi- cales qui se reproduisent soit en forêt soit en savane. Ce der- nier biotope traverse une période de disette à la fin de la sai- son sèche. En fait c’est vrai pour 6 des 8 familles de Passereaux africains dont on connaît le chiffre des pontes pour au moins 5 espèces dans chaque biotope. Il semble qu’il en soit de même pour 4 autres familles représentées par un nombre moins important d'espèces dans chacun des deux habitats ainsi que pour 2 familles tropicales américaines. 3°) Cette différence n’exisle pas entre les espèces de forêt et de savane chez les Muscicapidés et les Nectariniidés. 4°) Dans chacune des 4 familles, 2 d'Afrique et 2 d'Amé- rique, la moyenne du nombre d'œufs par ponte est plus forte en zone aride que dans les zones au climat moins rigoureux quoique non forestier. REFERENCES Ancner, G. F., and Gopmaw, E. M. 1961. — ‘The Birds of British Soma- liland and the Gulf of Aden, vols. 3, 4 (Edinburgh & London). Asxmoze, N. P. 1961. — The biology of certain terns. D. Phil. thesis (Oxford University). Cæapin, J. P. 1953, 1954. — The Birds of the Belgian Congo Parts 3, 4. Bull. Amer. Mus. Nat. Hist., 75, À, B. KLuuvEr, H, N. 1951. — The population ecology of the Great Tit. Ardea, 39 : 1-135. Lacx, D. 1947. — The significance of clutch-size. Ibis, 89 : 302-352. Lack, D. 1954. — The Natural Regulation of Animal Numbers (Oxford). Source : MNHN. Paris CLUTCH-SIZE IN TROPICAL PASSERINE BIRDS 89 Lack, D. 1958. — À quantitative brecding study of British tits. Ardea, 46 : 91-124. MancmanT, S. 1960. — The brecding of some S. W. Ecuadorian birds. Ibis, 102 : 349-382, 584-599. Monez, G, & Bouniiène, F. 1962. — Relations écologiques des avifaunes sédentaires et migratrices dans une savane sahélienne du bas Sénégal. Terre et Vie, 4 : 371-393. Monxau, R. E. 1944. — Clutch-size : a comparative study, with special reference to African birds. Ibis, 86 : 286-347. Permis, C. M. 1963. — Survival in the Great Tit, Parus major. Pro- ceedings XIIth International Ornithological Congress, pp. 717- Praëp, C. W. M., and Grant, C. H. B. 1955. — Birds of Eastern and North Eastern Africa, vol. 2 (London). Skuron, A. F. 1949. — Do tropical birds rear as many young as they can nourish ? Ibis, 91 : 430-455. Skurcu, A. F. 1954. — Life Histories of Central American Birds. Cooper Orn. Soc, Pacific Coast Avifauna, 31 : 19-121. Sxurcm, A. F. 1960. — Life histories of Central American Birds. Vol, 2 Cooper Orn. Soc. Pacifie Coast Avifauna, 34 : 287-577. WaGner, H. O. 1957. — Variation in clutch-size at different latitudes. Auk, 74 : 243-250. Wacwer, H. O. 1960. — Beziehungen zwischen Umweltfaktoren und der Brutzeit, Zahl der Gelege sowie ihrer Grôsse, Zool. Anz., 164 : 161-172. Source : MNHN. Paris CLASSIFICATION, IDENTIFICICATION AND SEQUENCE OF GENERA AND SPECIES by Ernst MAyR Museum of Comparative Zoology, Cambridge, U.S. A. Président du XIII Congrès International d'Ornithologie (Ithaca 1962) Much has been written in recent years about zoological classification. What is the purpose of classifying ? How should the findings of the taxonomist be presented ? In what sequence should species, genera, and families be listed in faunal lists and catalogues ? Regrettable as this is, it seems as if every generation has to face this problem anew. The fact that various kinds of organisms are clustered in nalural groups, was known long before scientific classifi- cations existed. One does not need to be a zoologist to know that there are birds and butterflies, hawks and ducks. Already the Greek philosophers speculated on the reasons for the existence of « natural groups ». In the 18th century the ques- tion of classification was foremost in the minds of the natu- ralists. The Aristotelians and their followers, including Lin- NAEUS, thought that taxa were natural because they reflected an underlying essence (the ideas of PLaro) which had rea- lity. The nominalists, on the other hand, with whom BUFFON and ADaANson sympathized, insisted that only individuals had reality, and that taxa were merely the creation of Man’s orde- ring mind. Neither interpretation was particularly satisfying, and in the 100 years after 1758 practising zoologists refused to be burdened unduly by philosophical speculations. Ins- tead, they classified organisms into the groups which seemed readily distinguishable by combinations of obvious charac- teristics. CUVIER’s recognition of four « embranchements » was one of the most important steps forward in the classifi- cation of animals. Many other zoologists made contributions to zoological classification that were hardly less important. The category of the family (unknown to LINNAEUS) was intro- duced into taxonomy around 1800 and the number of higher L'Oiseau et R.F.O., V. 35, 1965, n° spécial. Source : MNHN. Paris CLASSIFICATION OF GENERA AND SPECIES 91 taxa, from protozoans to mammals, recognized by zoologists rose rapidly. Students were attempting to design a « natural system », that is to recognize taxa which consisted of species resembling each other in the greatest possible number of cha- racters. They had no theory, however, that would explain why it was possible to construct a hierarchy of taxa, as is the natural system. It was not until after 1859 that general agreement was reached on the meaning of the higher taxa : Organisms fall into natural groups when they are the descendants of a com- mon ancestor, This concept gave new meaning to taxonomic activity ; it provided a working hypothesis to the taxonomist, who would ask himself « do the similarities of the organisms which I place in a single group indicate that they descended from a common ancestor ? » The theory'of classification now had a solid basis. This by no means resolved all the difficulties. One reason for this is that classifying has two rather different objectives. One is purely practical. A classification attempts to provide us with an arrangement to facilitate the identification of spe- cimens and their convenient sorting and storage in museum collections. As important as this practical aspect is, it is nei- ther the only nor, perhaps, the most important one. The other aspect is that a classification must have an explanatory value. Every new classification is, so to speak, a new taxonomic theory. Since we know that natural groups exist in nature only because all organisms are the products of evolution, it is our task, as scientists, to find what these groups are. It is our obligation to propose classifications which incorporate our researches on the most natural grouping of species and on the delimitation of such groups. Such classifications are of the nature of working hypotheses and are likely to be super- seded when better information becomes available. In this res- pect our work is in no way different from that in other branches of science. Nothing in science is ever final, nothing is every definitive, Every new scientific theory is an attempt to improve our understanding of nature, but it must be tes- ted against new observations again and again, and must be revised if it fails to explain the new observations. This is as true for every classification as it is for other scientific theo- ries. If new characters show that a previously proposed clas- sification is artificial, if it shows that a given species is pla- Source : MNHN. Paris 92 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE ced in the wrong species group or genus, then we must revise our classification in order to restore concordance with the new facts. This places us in a dilemma. For practical purposes we would like a completely stable classification, yet as scientists we cannot ignore improvements in our understanding of rela- tionships. What shall we do ? But this is not our only trouble. Evolution produces evolutionary trees, a multi-dimensional phenomenon, while a classification is always a linear sequence, a one-dimensional phenomenon. Even if we had a perfect understanding of evolution (which we do not have even in the best know groups), we would nevertheless be for- ced to adopt a linear sequence in which by necessity unrela- ted species are often placed next to each other. Yet, there is no other way of printing a list of species, or arranging spe- cimens in museum trays, than a linear sequence. Any list is a reference work, yet nothing is more annoying than to find that each of 3 or 4 lists one needs to consult, has adopted à different sequence. As Moreau (1961) points out quite correctly, it is difficult and time consuming to find a given species in such lists if every author adopts a different sequence. There are now some 5 or 6 different sequences of Passerine families in use at the present time. Some of these were proposed without the slightest published justification for the deviation from the last preceding list. This approaches scientific anarchy. What can be done to straighten out this chaos ? MOREAU’S suggestion has the unquestioned advantage of simplicity : Let us abandon all attempts to arrange species and genera accor- ding to presumed relationships, he proposes, but list them instead alphabetically, as was done by GESNER and TURNER 400 years ago. If there is so much uncertainty about rela- tionship, as is indicated by the differences of opinion, why not give up all pretense of trying to find a natural classifi- cation, he says, and adopt instead the simple and objective method of listing species and genera alphabetically ? The more so, since in many cases we have no information what- soever that would permit us to determine for a given species, genus, or family what its nearest relative is. Even though alphabetical lists of species are virtually unknown in ornithology, they are widespread in other branches of zoology, although usually only in name cata- Source : MNHN. Paris CLASSIFICATION OF GENERA AND SPECIES 93 logues rather than in monographic revisions. In those groups of insects, in particular, where a single genus may contain hundreds if not thousands of species, and where no good revision is as yet available, such an alphabetic arrangement is sometimes a necessity. But are the failings of natural clas- sification in ornithology really so grave that it would be advantageous to go back to the alphabetical system which ornithologists had given up 400 years ago ? To me it seems that an alphabetical system has far more short-comings than advantages, Let me mention some of these disadvantages. In collections il would place next to each other species that are neither similar nor closely related. Every time the name of a species is changed for nomencla- tural reasons, one would have to place it in a very different museum drawer, according to the first letter of its now valid species name. Every time the rank of a taxon is changed, as when a species is reduced to the rank of a subspecies, or a subspecies raised to that of a species, such a taxon would have to be shifted in the collections. The sequence of species in faunal lists would have to be changed for the same reasons. This would inevilably cause all sorts of inconveniences. Far more important is {he fact that an alphabetical arrangement lacks the great heuristic value of a natural — evolutionary) arrangement. By demanding that the most closely related species be placed next to each other, the theory of a natural arrangement continues to pose the ques- tion of relationship. Is species b so similar to species a, perhaps because it is only a subspecies of a, rather than a good species ? Are species c, d, and e, which are so similar lo each other, perhaps members of a single superspecies ? Is species which looks so different from species a to à perhaps a member of a different genus ? The requirement for placing species into natural groups does not permit the intellectually lazy solution of an alphabetical sequence. It demands deci- sions, it demands continuous study. No modern study of spe- ciation can be undertaken, unless based on a natural grou- ping of species. The splendid analysis of species in the genus Francolinus (HALL, 1963), for instance, could not have been undertaken without the prior sorting of the species into natu- ral groups. The same is true for Keasr’s (1961) analysis of the birds of Australia and all other modern speciation stu- dies in birds. They are all based on a natural grouping of species. Source : MNHN. Paris 94 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE There is one other very important reason for the scientific prestige of « natural » arrangements. The more we utilize taxonomic characters that can not be found in bird skins, but require the study of the living bird, his song, courtship habits, his biochemical constituents etc., the more important it becomes to test these new findings against those of mor- phological museum research. An alphabetical sequence of species would be of little use to someone doing comparative studies of serum proteins or of homologous bird calls. For all these reasons it seems to me that the traditional endeavor to discover and delimit natural groups, groups of related spe- cies, is still by far the best approach in classification. If taxo- nomy wants to maintain its role as a legitimate branch of biological science, it cannot abandon the endeavor to try to improve its theories, its classifications. Yet, this brings it at once into immediate conflict with one of the functions of classification, ease of reference. This function of a classifica- tion is, indeed, greatly weakened by the incessant changes in the sequence of species and genera. No taxonomist (including the writer of these lines) is innocent of this criticism. What can we do about it? Perhaps we should all agree not to publish any changes of standard classifications, of standard sequences (not necessarily the last published one !), until such changes have been formally adopted by a committee of the International Ornithological Congress. It is important to stress in this connection, as I have emphasized previously (Mayr, 1958), that a sequence is not a classification. The compromise then would be to adopt the principle of natural classifications, but not to permit every author to reshuffle the sequence of species, genera and families every time he has an « inspiration » that a different sequence would be « much better ». Even though the highest purpose of a classification is the explanatory one, we must never forget that classifications also have a practical purpose, and this is of course particularly true of sequences such as are adop- ted in faunal lists and in checklists. RESUME Tous les êtres organisés peuvent être classés en groupes naturels basés sur la filiation. Toute classification doit avoir à la fois : un rôle explica- Source : MNHN. Paris CLASSIFICATION OF GENERA AND SPECIES 95 tif, basé sur la filiation et la parenté, et un rôle pratique per- mettant l'identification et la mise en ordre des spécimens. Le rôle scientifique entre parfois en conflit avec le rôle pratique lorsque de nouvelles connaissances entraînent la modification de la classification préalablement admise. On cherche parfois à limiter le rôle de la classification à son objectif purement pratique en classant espèces et genres par ordre alphabétique. Ceci ne semble pas désirable pour des raisons non seule- ment scientifiques mais également pratiques. Les modifica- tions de nomenclature et de taxonomie entraînent des chan- gements même dans la séquence alphabétique. Le plus grand avantage du système alphabétique tient dans le fait qu’il n’a pas une valeur intrinsèque qui demande continuellement à être examinée et améliorée. Plus une classification demeure adoptée longtemps, mieux elle peut servir à des fins pratiques. Une classification ou une séquence taxonomique couramment acceptée ne devrait done être changée que lorsque de nouveaux faits viennent appor- ter des preuves de son caractère erroné. REFERENCES Haix, B. P., 1963. — The Francolins, A Study in Speciation. Bull. Brit. Mus. Nat. Hist., London, Zool., 10 : 107-204. Keasr, A., 1961. — Bird Speciation on the Australian Continent. Bull. Mus. Comp. Zool., Harvard College, Cambridge, 123 : 305-495. Mayr, E., 1958. — The Sequence of the Songbird Families. The Condor, 60 : 194-195. Moneau, R, E:, 1961. — Taxonomic Realism. Proc. Zool. Soc. Lond., 137 : 623-626. Source : MNHN. Paris ZUR TAXONOMIE VON CINCLUS CINCLUS EIN TIERGEOGRAPHISCHES PROBLEM von Günther NIETHAMMER Museum Alexander Koenig, Bonn Zwischen Nord- und Südspanien kônnen bedeutende tier- geographische Unterschiede festgestellt werden, die dem sehr unterschiedlichen Klima zwischen beiden Teilen der Iberi- schen Halbinsel entsprechen. Das nôrdliche Küstenland Spa- niens gehôrt zu den regenreichsten Gebieten Europas, Mittel- und Südspanien hingegen sind relativ trocken. Die nordspa- nischen Gebirge bilden als natürliche Fortsetzung der Pyrenäen nach Westen mit diesen zusammen eine zoogeogra- phische Einheit. Das kommt zum Ausdruck durch eine Anzahl eurosibirischer Arten aus vielen Tierklassen, die in den Pyre- näen und nordspanischen Gebirgen, nicht aber sonst auf der Iberischen Halbinsel vorkommen. Beispiele hierfür liefern unter den Säugetieren Rupicapra rupicapra, Sorex minu- tus (*), Neomys fodiens (*), Micromys minutus (*), Clethrio- nomys glareolus (*), Arvicola terrestris (*), Microtus agres- tis (*) und Apodemus flavicollis (*), unter den Vôgeln Tetrao urogallus, Scolopax rusticola, Dryocopus martius, Anthus tri- vialis, Lanius collurio, Locustella naevia, Phylloscopus tro- chilus, Saxicola rubetra, Turdus philomelos, Parus palustris und Pyrrhula purrhula. Etliche auf Nordspanien beschränkte Säugetiere haben im Süden einen naheverwandten Repräsen- tanten wie Arvicola terrestris — A. sapidus, Talpa europaea — T. caeca, Microtus arvalis — M. cabrerae und Neomys fodiens — M. anomalus. Diese Unterschiede im Artenbestand werden noch deut- licher gemacht durch den Umstand, dass manche auf der Iberischen Halbinsel weitverbreitete Arten im N und S$ verschiedene Subspecies ausgebildet haben. Bei vielen Klein- säugern ist eine Verdunkelung des Felles entsprechend den (*) Dieses Arten haben wir 1963 zum ersten Male in Nordspanien gefunden (J. NirmamMEr, Z. Säugetierkunde, 1964, pp. 193-220). L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris TAXONOMIE VON CINCLUS CINCLUS 97 hohen Niederschlägen in Nordspanien offensichtlich, bei den Vôgeln ist dies nicht nachgewiesen oder doch weniger deutlich. Ein schônes Beispiel für eine klare rassische Dif- ferenzierung zwischen den Populationen von N und S geben die beiden Unterarten der Schwanzmeise, Aegithalos caudatus taiti und irbü, die VAURIE sogar zwei verschiedenen Rassen- gruppen zuteilt. Andere Beïspiele zeigen eine nur schwache Differenzierung der Populationen wie etwa (die in N-Spanien lebende Rasse jeweils zuerst genannt) Oenanthe oenanthe oenanthe und nivea, Sitta europaea caesia und hispaniensis, Parus caeruleus caeruleus und ogliastrae sowie Parus ater aler, vieirae und cabrerae. Es gibt unter den Vôgeln auch Arten, deren iberische Unterart über die ganze Halbinsel von $ bis N verbreitet ist. Beispiele hierfür sind Phylloscopus collybita brehmi und Phoenicurus ochruros aterrimus. Diese letztere Rasse gehôrt zu jener nicht sehr zahlreichen Gruppe von Unterarten, die im Felde von ihren geographischen Ver- tretern sicher unterschieden werden kônnen wie etwa Nebel- und Rabenkrähe, Haus- und Italiensperling. Ich habe in den Karsthergen von Ramales de la Victoria in Nordspanien im April 1963 Männchen des Hausrotschwanzes gesehen und wiederholt eingehend beobachtet, die einen kohlschwarzen Rücken und Scheitel hatten und darin soger dem dunklen Extrem von Ph. o. aterrimus glichen. Ein deutlich gekennzeichnetes Rassenpaar wurde in Spa- nien dagegen bei der Wasseramsel konstatiert. Als erster hat dies WrrmenBy (1922, 1928) klar erkannt. 1957 pflichtete ich ibm auf Grund neuen Materials aus Süd- und Nordspanien bei, wogegen VAURIE zunächst (1955) auf der ganzen Ibe- rischenHalbinsel nur eine Rasse der Wasseramsel anerkennen wollte. 1958 revidierte er aber seine Ansicht und schloss sich gemeinsam mit GREENWAY der Auffassung WirHERBys an. Ich hatte 1963 und 1964 Gelegenheit, Wasseramseln in Nord-, Mittel- und Südspanien zu sammeln, nämlich 2 bei Ramales de la Victoria, 1 in der Sierra de Guadarrama (Südhang), 2 in der Sierra de Cazorla und 1 (durch Dr. Ags) in Linares bei Salamanca. Diese konnte ich mit dem schon im Museum Koenig, Bonn, vorhandenen Material (insbesondere aus Val- ladolid) und einer Serie aus den Pyrenäen, die mir Dr, Mac- DONALD freundlicherweise aus dem British Museum, London, lieh, vergleichen. Alle nôrdlich Madrids und in den Pyrenäen gesammelten Source : MNHN. Paris 98 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Stücke sind dunkelbäuchig, die südspanischen rostbäuchig und nicht von typischen aquaticus zu unterscheiden. Ganz Nordspanien und die Pyrenäen nimmt also eine dun- kelbäuchige Form vom cinclus-Typ ein, die Gebirge Südspa- niens eine rostbäuchige Form vom aquaticus-Typ. Das Ver- blüffende hierbei ist, dass die dunkelbäuchigen spanischen Wasseramseln nicht oder kaum von skandinavischen C. c. cinclus, die rostbäuchigen nicht von mitteldeutschen C. c, aqualicus zu unterscheiden sind, obwohl die jeweils ähnlichen bzw. gleichen Populationen (Nordeuropa — Nordspanien, Mitteldeutschland — Südspanien) keine Verbindung mitei- nander haben. Wie ist dieses Verbreitungsbild entstanden ? Es gibt zwei Môglichkeïten es zu deuten, nämlich — ükolo- gisch — als Ergebnis konvergenter Entstehung jeweils zweier gleicher Rassen in den heutigen Arealen oder — historisch — als Ergebnis einer Arealaufspallung zweiïer einst kontinuier- lich verbreiteter Rassen. Die erste Deutung würde gestützt, wenn die ükologischen Bedingungen, unier denen jeweils gleichgefärbte Populationen leben, gleich oder ähnlich wären. Das ist offensichilich nicht der Fall : für die Alpen, Pyrenäen und Kantabrischen Gebirge sind z. B. die klimatischen Fak- toren äbnlich, aber wir finden hier 2 Rassen ; in den Alpen und Gebirgen Südspaniens ist das Klima unterschiedlicher, aber die Wasseramseln sind hier wie da rostbäuchig. Aus der heutigen Verteilung der beiden Rassen bzw. Ras- sengruppen (sowohl unter den dunkelbäuchigen wie unter den rosthäuchigen Populationen sind jeweils mehrere recht schwach unterschiedene Formen beschrieben worden) ist weder ersichilich, wo sie entstanden und ursprünglich ver- breitet waren, noch welche Ausbreitungs- oder Rückzugswege sie genommen haben. Auffallend ist nur, dass Kleinasien genauso wie Iberien von beiden bewohnt wird (1), und zwar jeweils von cénclus-artigen im Norden und aquaticus- artigen im Süden, So ist C. c. cinclus auf 3 weit voneinan- der isolierte Areale verteilt, nämlich erstens Skandinavien samt Baltikum bis Ostpreussen, zweitens Nordostkleinasien und drittens Pyrenäen, Nordspanien und die beiden Inseln Korsika und Sardinien. Zwischen diese 3 Bereiche erstreckt sich als breite Zone von Schottland und Irland bis Peloponnes @) C. c. amphytrion (aus Lasistan) ist nicht von cinclus zu unterschei- den. Dagegen gehôren Wasseramseln aus dem Taurus zur uquaticus- Gruppe (KumerLorve, 1961). Source : MNHN. Paris TAXONOMIE VON CINCLUS CINCLUS 99 und Sizilien das Areal von C. c. aquaticus oder aquaticus- äbnlichen Populationen. Die Verbindung nach Südspanien ist über Sizilien und NW-Afrika, nach Südanatolien über Pelo- ponnes und Cypern gegeben. Im Libanon, Kaukasus und in Persien schliessen sich weitere rosthäuchige Formen an und im Ural lebt weithin isoliert C. c. uralensis, der zwischen cinclus und aquaticus ungefähr in der Mitte steht. Dieses Ver- breitungsmuster der Cinclus-Rassen bzw. der beiden Farb- typen in der westlichen Paläarktis historisch erklären zu wol- len, etwa als Folge postglazialer Arealverschiebungen zweier durch die Eiszeit getrennter und während dieser entstande- nen Rassen, scheint mir sehr schwierig, wenn nicht unmôüg- lich zu sein. Es bleibt demnach kaum eine andere Erklärung als die Annahme, dass die dunkelbäuchige Form der Wasser- amsel an 3 Stellen unabhängig voneinander entstanden ist (G. c. cinclus in Nordeuropa, C. c. pyrenaicus in N-Spanien und C. c. amphytrion in N-Kleinasien). Ich habe bisher die klare Trennung von dunkel- und rostbäuchigen Populationen betont und insbesondere in Spanien weder bei der einen noch bei der anderen Form Ausnahmen gefunden. Die Populatio- nen der Wasseramsel, insbesondere diejenigen des aquaticus- Typs, zeigen aber auch in diesem so auffälligen Merkmal eine erhebliche individuelle Variation, die von Ricnrer (1954), BALAT (1961) und Josr (briefl.) (1) untersucht worden ist. Ricmrer fand bei 65 sächsischen Vôgeln 9 dunkelbäuchige, BaLar bei 79 tschechischen 3 und Josr bei 56 hessischen keine dunkelbäuchige ; mit anderen Worten : unter 200 G. c. aquaticus waren 12, d. h. 6 %, vom cinclus-Typ. Das bedeutet, dass in den Populationen von C. c. aquaticus auch die cinclus-Mutante « griffbereit » vorliegl, die sich anrei- chern müsste, sobald ein entsprechender Auslesefaktor wirk- sam wird. Eine Parallelentwicklung von geographischen Rassen einer Art, wie sie uns als Ergebnis der gleichen auslesenden Fakto- ren z. B. bei den afrikanischen Lerchen bekannt ist, kônnte durchaus auch für die disjunkte Verbreitung von C. c. cinclus verantwortlich sein. Dass man bisher (s. 0.) eine Korrelation zwischen der Bauchfärbung und klimatischen Faktoren nicht feststellen konnte, ist kein Grund, die Wirkung der Auslese () Herrn Otto Josr, Fulda, der gefangene Wasseramseln jeweils unmittelbar mit Bälgen verschiedener Typen verglich, bin ich für Mit- teilung dieses unverôffentlichten Materials dankbar. Source : MNHN. Paris 100 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE zu verneinen, denn die Bauchfärbung kann ja genetisch mit einem anderen, der lokalen Auslese unterworfenen Merkmal gekoppelt sein. RESUME L'étude des différences biogéographiques entre le Nord et le Sud de la Péninsule ibérique a ‘amené l’auteur à se pencher sur le cas du Cincle, qui dans le Nord de l'Espagne et les Pyrénées se présente sous le type cinclus à ventre pâle, et dans le Sud sous le type aquaticus à ventre couleur rouille. Les oiseaux appartenant à ces deux formes ne se distinguent pratiquement pas de ceux de Scandinavie (cinclus) ou d’Alle- magne (aquaticus). L'auteur discute la taxonomie de l’espèce, n’adoptant pas l’idée de convergences écologiques mais plu- tôt celle d’une évolution parallèle des races dans une aire discontinue. LITERATUR BaLat, F. (1961). — Colour Variability in the Dipper, Cinclus cinelus (L.), in Central Moravia and its Taxonomic Relations. Zoologicke Listy, X, p. 135. Bennis, F. (1954). — Prontuario de la Avifauna. Ardeola, 1, p. 60. Gneexway, J. C, & Ch. Vaurte (1958). — Remarks on some forms of Cinelus (Aves). Breviora, Nr. 89, pp. 1-20. KUMERLOEVE, H. (1961). — Zur Kenntnis der Avifauna Kleinasiens. Bonn. zool. Beitr., 12, Sonderheft, pp. 134-135. NIETHAMMER, G. (1957). — Zur Vogelwelt der Sierra Nevada. Bonn. zool. Beïtr., 8, pp. 241-242. Ricuren, H. (1945). — Zur Variabilität der Färbung sächsischer Wasser- amseln. J. Orn., 95, p. 54. Vaum®, Ch. (1951). — Notes on the Wrens and Dippers of Western Asia and India. Am. Mus. Nov., 1485, pp. 9-15. — (1955). — Systematic Notes on Palearctic Birds. No. 16. Troglody- tinae, Cinclidae, and Prunellidae. Am. Mus. Nov. 1751, pp. 12-11. — (1959). — Birds of Palearetie Fauna. Passeriformes. London, pp. 193-198. Source : MNHN. Paris LE MARTINET PALE DE SOCOTRA (APUS PALLIDUS BERLIOZI) by S. Diccon RIPLEY Secretary, Simthsonian Institution During the past winter the U. S. National Museum was so fortunate as to obtain permission for Mr. Alec FORBES- WATSON to visit the remote island of Soqotra east of Cape Gardefui (Lat. 12°30° N., Long. 54° E.). The kindness of the Sultan of Qishin and Soqotra in granting this permission is most gratefully acknowledged. Among the specimens taken by Mr. FoRBEs-WATsoNn is a series of pale swifts. OGiLvie-GRANT, in FORBEs (1903) recor- ded seeing the species but no one had ever collected it. I wish now to dedicate this swift to my distinguished colleague and friend of many years, on the occasion of this honorary issue of L’Oiseau et la R.F. 0. : Apus pallidus Berliozi, new subspecies Type : g USNM No. 518025, collected by Mr. Alec For8Ees-WaTson at Kishin, Hagghier Mts., Soqotra, altitude 2200 feet, on 3 May 1964. Description : From somalicus (Clarke), 1919, this form differs by being larger and darker, especially on the under- parts which are richly blackish rather than fuscus. From brehmorum of the western Mediterranean this form differs by being more grayish-blackish above and below, less brownish or olive-brown tinted, and by being smaller. Measurements : wing tail Berliozi 20 Gé 163.5-178 (169.8) 67.5-77 (70.9) mm 1299 155-172 (166.5) 67-70 (68.8) somalieus 639 152-156 (154.2) 65 Brehmorum 7149 164-182 (175.6) 70-80 (77) L'Oiseau et R.F.O., V. 35, 1965, n° spécial. Source : MNHN. Paris 102 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Weight : Berliozi Go 34-438 (38.5 + 4.5) gms; 9 Q 37- 46 (+ 4.5) gm. Color of soft parts ; iris dark brown, bill black, feet dark pink-brown, or brown-maroon, claws black. Range : Soqotra Island, western Indian Ocean. These birds were nearly all in breeding condition, some of the females containing large yolks. They were collected between April 14 and May 9, 1964, most of them from 2000- 3800 feet in the mountains. Additionally two males were taken May 17 at sea level one of which was not in breeding condition. Mr. ForBes-Warson felt that the breeding season presumably was virtually over by mid-May. He noted stomach contents as black beetle or flying ant fragments, and in two cases, one at sea level, grasshoppers. On May 18 he watched swifts catching flying termites over Hadibu town at sea level after a rain shower (1). RESUME Description d’une nouvelle sous-espèce de Martinet pâle collecté par Alec ForBes-WaTson à Socotora en 1964, et que l’auteur dédie au Professeur BERLIOZ à l’occasion de ce numéro jubilaire. Depuis cette époque, une nouvelle sous-espèce a été décrite par FRIEDMANN en Ouganda, mais celle-ci diffère nettement de la race Berliozi. (1) Since writing the above, D' Herbert Frepmanx has described Apus pallidus kapnodes (1964), Los Angeles County Museum Contributions in Science, N° 83, Dec. 8) from Mt. Moroto, Uganda. Through D° Friep- Man's kindness I have been able to examine six specimens which con- firm that Berliozi differs significantly from Kapnodes in larger size (apnodes ; wing d 151-158 [154.21 ; © 151-160 [156]), as well as in plumage pattern. In kapnodes the white throat is sullied by an infusion of brownish along the dark shaft and adjacent vanes of the basal and median parts of the featers. This series of kapnodes is also far darker below and somewhat darker above than Berliozi, with greatly reduced white tips or edgings to the breast and abdomen feathers. Source : MNHN. Paris NOTES ON THE SUNBIRD-ASITYS (NEODREPANIS) by Finn SALOMONSEN Zoological Museum, Copenhagen This volume ïis dedicated to Prof. Jacques BErLio7, and I find it appropriate to pay him a hommage by commen- ting on some exotic and little known birds. Prof. BERLIoZ, whom I have known for almost 40 years, has always been particularly fond of puzzling and exciting species inhabiting remote areas, and he is one of the very few ornithologists of our days that are equally well acquainted with the fauna of all five continents. His experience concerning avian systema- ties is enormous, and I have always admired his world-wide knowledge as well as his sound and conservative view-points in taxonomic matters. I take this opportunity to thank him for many stimulating discussions in correspondance and in personal interviews in Paris, in Copenhagen and at interna- tional conferences abroad. The bird life of Madagascar is only one of Prof. BERL107” many interests, and he may now recall that many years ago I called a very distinct Malagasy bird after him (Mento- crex kioloïdes berliozi, SALOMONSEN 1934 a, p. 386) as a token of my gratitude for his indefatigable help during my work with the Madagascar avifauna. The following article is only a rather short note, but it deals with some very interesting Malagasy birds. When Neodrepanis coruscans was discovered by CROSSLEY in eastern Madagascar and described by SHARPE in 1875 it was by HAaRTLAUB (1877, p. 94) called « one of the most inte- resting ornithological discoveries in recent time ». 1 still find that it represents a most interesting little group of birds. The genera Philepilta and Neodrepanis, each with two spe- cies, form the endemic Malagasy family of the asitys (Phile- pittidae). Whereas the two Philepitta are short-billed fruit- L'Oiseau et R.F.0., V. 85, 1965, n° spécial. Source - MNHN. Paris 104 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE eating species, the Neodrepanis have a long and very thin, strongly curved bill, reminding of that in certain sunbirds, such as Drepanorhynchus, honey-eaters such as Acantho- rhynchus and drepanidids such as Hemignathus and others. In all these groups a convergent evolution has taken place, due to similar adaptations. The Neodrepanis have all ear-marks of being nectar- feeding birds, and the scanty information on their life-habits supports this assumption. RanD (1936, p. 472) described how N. coruscans continually came to a flowering tree, and during two days sixteen specimens were collected there, and just as many were observed, while in the forest elsewhere only a single individual was recorded. MiLNE-EDWARDS and GRANDI- piEr (1879, p. 290) mentioned that N. coruscans is often found at the balsam species Impatiens Humblotiana, which is widely distributed in the rain-forests of Eastern Madagascar. This plant has large red blossoms, with the corolla forming a long eurved tubular spur, almost of the same size and form as the bill of Neodrepanis. This structure is undoubtedly an adap- tation to ensure pollination through visits by Neodrepanis, and is analogous to similar floral modifications in the ende- mic ornithophilous Lobeliaceae of the Hawaiian Islands, which are adapted to pollination by drepanidids. On the colour-plate accompanying my paper on Neodrepanis (SALo- MONSEN 1934) these birds are seen feeding in a flowering Impatiens Humblotiana. Not only the bill, but even the tongue demonstrates adap- tations to nectar-feeding. As shown by AmaDON (1951, p. 62) the tongue of Neodrepanis is tubular, but differing in struc- ture from that in other nectar-feeding passerines, while the tongue of Philepitta is unspecialized. The remarkable structural differences between Philepitta and Neodrepanis illustrate an adaptive radiation in this family which, it is true, is by far not so striking as the well- known cases in Geospizinae, Drepanididae and Vangidae, but nevertheless is an interesting phenomenon. Neodrepanis has been able to utilize an ecological niche which was virtually unoccupied in Madagascar. Only the long-billed Cinnyris notatus may occasionally compete with it during feeding in flowers, as noticed already by RanD (loc. cit.), while Cinnyris sovimanga and Zosterops madaraspatanus have much too short bills. Source : MNHN. Paris NOTES ON THE SUNBIRD-ASITYS 105 The English vernacular of Neodrepanis was formerly < Wattled Sunbird », but when it was discovered by AMADON (951, p. 59) that these birds belonged to the Philepittidae the name was changed to « False Sunbird » (e.g. by GiLLrarD 1958, p. 291, and Rann 1964, p. 66), following Amapon’s French designalion « Pseudo-Souimanga ». It appears to me, however, Lo be more appropriate to call these birds « Sunbird- Asitys », in acknowledgement of their position as members of the family of asitys. It is peculiar that the close relation- ship between Neodrepanis and Philepitta was discovered so late, although a number of expert ornithologists, such as SHARPE, HARTLAUB, GRANDIDIER and SHELLEY have studied these birds. Some of these students, however, had a feeling that the two genera were related, this holding good especially of SHELLEY. The similarities between Neodrepanis and Philepitta include the following characters : (1) Plumage coloration, particularly noticeable in males of Philepitta schlegeli and Neodrepanis hypoxantha, which both have the under-parts uniform yellow of about the same shade ; (2) Sexual dimor- phism, with greenish plumage in females ; (3) Very short tail, constituting about half the wing-length (in sunbirds usually at least two-thirds the wing-length) ; (4) Presence of orbital wattles in adult males (not developed in sunbirds) ; (G) Extremely long first (outer) primary, extending almost to tip of wing (in sunbirds constantly shorter than secondaries). To this come the important and decisive family characters, demonstrated by AMADoON (loc. cit.) : (6) Mesomyodian syrinx, (7) Taxaspidian scutellation on tarsus, (8) Vestigial hyporha- chis (well-developed in sunbirds). When I studied these birds it did not appear to me, admit- tedly, that Neodrepanis and Philepitta were related, even though in my paper (SALOMONSEN 1934, p. 2) I pointed out and discussed the extraordinary length of the first primary in Neodrepanis, which was unprecedented in the suborder Oscines. It may be some excuse for me that I was mainly con- cerned with the study of the moult and sequence of plumages, in which respect the situation is quite extraordinary. The moult is extremely similar in Neodrepanis and the sunbirds, while it differs strikingly in Philepitta. The members of the latter genus has only one annual, postnuptial moult, while Néodrepanis has two annual moults, developing a special off- Source : MNHN. Paris 106 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE season (eclipse) plumage, which is the case also in most sun- birds. The parallelism goes so far that even the extent of the partial, praenuptial moult in Neodrepanis equals that in cer- tain sunbirds. This is a peculiar situation. During a visit to the British Museum (Nat. Hist.) I stu- died the moult of Neodrepanis coruscans, and found that the similarity with that in the Mascarene sunbirds, especially Cinnyris sovimanga, was striking. No specimens of N. corus- cans from the period October-March were examined, but the September birds were in breeding plumage and had big and swollen testes. The breeding-time, therefore, is undoubtedly September-November, as it is also in Cinnyris sovimanga, and for that matter in the greater part of the birds inhabiting Eastern Madagascar. I have seen no nestlings (in natal plumage) of N. corus- cans, but there are many immature birds (in juvenal plu- mage) in the collection of the British Museum. The juvenile males and females are indistinguishable, and I also failed to see any difference between the juvenile birds and the adult females. The juvenal plumage is kept to May-July, when it is replaced through a complete postjuvenal moult by the first nuptial plumage, in which the birds very likely already breed. This corresponds with the situation in Cinnyris notatus in which species the juvenal plumage is shed through a complete postjuvenal moult in May-July. Cinnyris sovi- manga has, on the other hand, a complete postnatal moult, and the juvenile birds are quite like the adult birds in off- season dress. The adult males of N. coruscans have, as already men- tioned, alternating off-season and nuptial (breeding) plu- mages. This siluation is present also in most sunbirds, and in the Mascarene species I have found it in all except Cin- nyris notatus (and its subspecies) and C. dussumieri, which have only one moult. Adult specimens of N. coruscans col- lected in April are still in full off-season plumage, but already some May birds (males) are in breeding plumage, but some moult later, as shown by a specimen captured in August and still in its prenuptial moult. This corresponds with the con- dition in Cinnyris sovimanga, of which a large series was examined. In this species the prenuptial moult extends from June to August. Neodrepanis, just as all other passerine birds, has a com- Source : MNHN. Paris NOTES ON THE SUNBIRD-ASITYS 107 plete postnuptial moult, including wings and tail, but owing to lack of material from October-March I have seen no spe- cimens in postnuptial moult. In the Mascarene sunbirds 1 have found the following periods for the complete post- nuptial moult : C. sovimanga : January-March, C. coquerelli, C. aldabrensis, G. abbotti and C. dussumieri : March-April. As the time for the partial prenuptial moult of Neodrepanis corresponds with that in C. sovimanga it is probable that its complete postnuptial moult takes place in January-March, just as in C. sovimanga, and this would signify that Neodre- panis kept its off-season plumage for 4-5 months, from January-March to May-August. À very interesting specimen of N. coruscans is a male in full prenuptial moult, captured on the 23. August 1930 at Andapa. The moult is incomplete, and the feathers are renewed in the following regions : Head, nape, hindneck, mantle, back, scapulars, ear-coverts, checks, chin, throat and upper breast, but not on rump, lower breast, abdomen, wing- coverts and upper and under tail-coverts and of course not remiges and rectrices. The rump, the tail-feathers and the smaller wing-coverts are already glossy in the off-season dress, and through the prenuplial moult also the front, cheeks, neck, mantle, back and scapulars attain metallic glossy feathers. It is noteworthy, however, that the new fea- thers on chin, throat and upper breast are olive-yellow with greyish bases without the least metallic gloss, ie. exactly as in the off-season plumage. Consequently Neodrepanis moults its feathers on chin, throat and upper breast twice annually, but the two feather gencrations are identical in coloration, In C. sovimanga the partial prenuptial moult comprises the same regions as in Neodrepanis, but the new feathers on throat and breast are bright metallie, while in the off-season dress they are yellowish with blackish bases, of the same colour as the feathers of the back. Just as in Neodrepanis the rump feathers in C. sovimanga are not renewed during the prenuptial moult, but contrary to the glossy-rumped Neo- drepanis they are brownish grey without gloss and remain so in the breeding plumage. The remarkable similarity in moult between Neodrepanis and the sunbirds is not easy to explain. We know now that it does not express any actual relationship, but is due to a parallel development. It is difficult to understand, however, Source : MNHN. Paris 108 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE how this parallelism could be the result of similar adapta- tions, when bearing in mind that the sunbirds, which all have very similar life-habits, demonstrate a considerable variation in their moulting schemes. The above-mentioned specimen of Neodrepanis coruscans (from the 23. August) demonstrates very well the prenuptial development of the orbital wattles. In the off-season dress the wattles are obliterated and the skin is completely feathe- red. In the specimen in question most of the off-season fea- thers are shed, but not renewed, with the result that the orbital region, 2-3 mm in front of the eye and 4-5 mm behind the eye, is almost bare, supplied only with scattered and very worn feathers belonging to the off-season dress. At the same time the skin has begun to arch backwards (caudally), and a lobe of about 1.5 mm in length has already developed. At a later stage the lobes grow still larger, and in the breeding time the bare skin extends up to 5 mm above the eye and twice as much behind the eye. After the breeding time the lobes disappear and the orbital region is again supplied with feathers. Some words should be added about the species Neodre- panis hypoxantha, which differs from N. coruscans by having, in both sexes, uniform bright canary-yellow under-parts (slightly lighter in female) and by having horn-brown (not black) bill, which is distinctly shorter and finer and not so cürved as in coruscans ; further, by having, in adult males, a much stronger emargination on the first (outer) primary, which has the tip very strongly attenuated, and, finally, by possessing a loral extension of the orbital wattle, This distinet species is much rarer than N. coruscans and, apparently, has a much more restricted range. Only very few specimens are known. I described this species (SaLo- MONSEN 1933, p. 182) on the basis of two specimens (adult male in off-season plumage and adult female) in the British Museum, collected by the Rev. W. Dean Cowan in July 1881 in the forests east of Tananarive. For some years these two specimens were the only ones known, but in 1937 STRESE- MANN announced that there were two specimens (adult male in breeding plumage and adult female) in the Zoological Museum in Berlin and one more adult male in another Ger- man museum, all three specimens collected in November 1880 by J. M. HipeBranDT at a locality called Andrangoloaka, Source - MNHN. Paris NOTES ON THE SUNBIRD-ASITYS 109 situated somewhat east of Antsirabé on the slopes of the plateau in central eastern Madagascar (STRESEMANN 1937, p. 135). Subsequently, Wermore (1953, p. 91) published a further record, an adult male in breeding plumage, kept in U. S. National Museum in Washington, collected by the Rev. James Wizzs in October-November 1895 in « E. Imerina », which is the name of the plateau around Tananarive, now almost devoid of forest. Until this period all specimens known, a total of six, were collected on the mountain slopes of the central eastern parts of Madagascar, and in these areas the forest has been vir- tually destroyed. More recent expeditions, such as the great Mission Zoologique Franco-Anglo-Américaine à Madagas- car 1929-30, did not meet with N. Aypoxantha at all, with the result that this species was generally supposed to be extinet, e.g. by the present author (SALOMONSEN 1934, p. 6), STRESEMANN (1937, p. 136) and Wermore (1953, p. 91), and even as recently as 1958 by GizciarD (1958, p. 291). In the same year, however, GREENWAY (1958, p. 28) mentioned a specimen (male) collected in the Sianaka forest the 95. February 1925 and now in the Museum of Comparative Zoo- logy in Cambridge, Massachusetts. The specimen was pur- chased from the dealer Karl FRiTSCHE in Bremerhaven, Germany, who in those years sold large numbers of Mada- gascar skins, collected in the Sianaka forest. This forest, situated between Fanovana and Lac Alaotra, is very exten- sive and not much destroyed, and GREENWAY coneludes that < N. hypoxantha will be found again ». This prediction has come true, a still more recently collected specimen has turned up. During a visit to the British Museum a few years ago I found in the collection an adult male in off-season plumage of N. hypoxantha, misidentified as N. coruscans, collected in August 1929 in the « Grande forêt de l'Est (Fito) » by the well-known French forester and ornithologist L. LAVAUDEN. The specimen is catalogued under the number 1939. 12. 9. 588. Fito is a locality situated within the great forest of Sia- naka. Although I have been informed that the destruction of the Sianaka forest has strongly increased in recent years I am quite certain that N. hypoxantha still survives in these areas, which is the main stronghold of many other rare species, such as Hartertula flavoviridis, Dromaeocercus brun- neus, Oxylabes cinereiceps, Oxylabes xanthophrys and Ber- Source : MNHN. Paris 110 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE nieria tenebrosa, and probably also of the very rare Newlonia fanovanae, of which the type specimen still is the only one known. I can add that I have located one more specimen of N. hypoxantha collected by J. M. HILDEBRANDT. It belongs to the collection of the late Count O. Zepzirz, and is now kept in the Riksmuseum, Stockholm. The specimen is an adult male in breeding plumage, collected in November 1880 at Andrangoloaka, just as the three other HiLDEBRANDT Spe- cimens. Of the nine known specimens of Neodrepanis hypoxantha seven are males, and there are information about the plu- mage in six of these. Four males collected in October- November are in the breeding plumage, and two collected in July-August are in the off-season plumage. In the July spe- cimen (the type of the species) a few feathers of the breeding plumage have already appeared on the mantle. Although the material is scanty it tends to show that the sequence of plu- mages in the adult male of N. hypoxantha is similar Lo that in N. coruscans. RESUME La famille malgache des Philepittidés est un exemple de radiation adaptative due, en grande partie, aux différences dans le comportement alimentaire des espèces appartenant aux genres Philepitta et Neodrepanis. La mue et les séquences de plumage de ces deux genres sont ici décrites. Les Neodre- paris, qui sont nectarivores, ressemblent superficiellement aux Souis-Mangas (Nectarinidés). Ils ont une mue qui, dans le détail, ressemble également beaucoup à celle des Souis- Mangas. Pourtant, les deux groupes sont sans relation et leur ressemblance est due à une simple évolution parallèle. La position systématique du rare Neodrepanis hypoxantha fait l’objet d'une discussion. Cette espèce a été considérée comme éteinte, mais des spécimens ont été collectés en 1925 et 1929 dans la forêt de Sianaka. Aussi l’auteur en conclut-il que l'espèce existe encore dans cette région où la forêt n'a pas été détruite. Source : MNHN. Paris NOTES ON THE SUNBIRD-ASITYS li REFERENCES Amapox, D., 1951, — L'Ois. et la R.F.0., vol. 21, pp. 59-63. Guzranp, E. Th., 1958. — Living Birds of the World. New York (400 p.). Greënway, Jr, J. C., 1958. — Extinct and Vanishing Birds of the World. New York (518 p). Harriaur, G. 1877. — Die Vôgel Madagascars und der benachbarten Insel- gruppen. Halle (426 p.). Mine-Enwanrps, A. et A. GRANDIDIER, 1879. — Histoire Physique, Natu- relle et Politique de Madagascar, vol. 12. Paris. Rax», A. L., 1936. — Bull. Amer. Mus. Nat, Hist., vol. 72, art. 5, pp. 143- 499. — 1964, — Asity, in A. LanpssorouG# Thomson (Ed.), A New Dic- tionary of Birds. London (928 p.). SaLoMonseN, F., 1933. — Bull, Brit. Ornith, Club, vol. 53, pp. 182-183. — 1934. — L'Ois. et la R.F.0., vol. 4 (N.S.), pp. 1-9. — 1934a. — The Ibis, ser. 13, vol. 4, pp. 382-390. Srresemann, E., 1937. — Ornith. Monatsber., vol. 45, pp. 135-136. Wermore, A. 1953. — The Auk, vol. 70, p. 91. Source - MNHN. Paris MOLECULAR SYSTEMATICS NEW TECHNIQUES APPLIED TO OLD PROBLEMS by Charles G. SIBLEY Yale University, New Haven, Conn., U.S.A. Secrétaire Général du XIII° Congrès International d'Ornithologie ({thaca 1962) The search for evidence of taxonomic relationships has prompted systematists to investigate many aspects of the form and function of birds. NirzscH (1840) produced one of the first broad comparative studies with his atlas of pterylo- graphy and MüLLer (1847) examined the structure of the syrinx in many groups of birds. In 1867 T. H. HuxLey deve- loped a classification based upon the palatal bones and a few years later GaRRoD made recommendations for classifying birds on the basis of the carotid arteries (1873 a) and the arrangement of the pelvic muscles (1873b ; 1874). Garron also (1875) found useful information in the flexor tendons and GAabow (1899) reported on the taxonomic value of the intestinal convolutions of birds. FORBES, FURBRINGER, BED- DARD, PYCRAFT, SHUFELDT and others studied various anato- mical characters including the structure of the toes, the con- dition of the fifth secondary (eutaxy vs diastataxy), the shape of the nostrils (schizorhinal vs holorhinal), the presence or absence of a septum between the nares, the arrangement of scutes on the tarsal envelope, the variations in the sternum and tongue, the size of the intestinal caeca, the development and feathering of the uropygial gland, the condition of the aftershaft and the thigh artery and the number of cervical vertebrae, remiges, rectrices and many other structures. In more recent years some of these have been re-examined and additional anatomical and functional characters have been investigated. Hupson (1937 ; 1948) has added to the work on pelvie muscles, GLENNY (1955) has studied the carotids and other arteries in the region of the heart, BEECHER (1950) compared the jaw muscles of the passerines, Torporr (1954) considered certain aspects of the skull of the Fringillidae L'Oiseau et R.F.O., V. 35, 1965, n° spécial. Source : MNHN. Paris MOLECULAR SYSTEMATICS 113 and ÂMES (in press) has studied the passerine syrinx. STALL- cup (1954 ; 1961) and Mararpi (1963) have used serology and several studies of behavior have made taxonomic recommen- dations. The parasites of birds have been used as evidence of avian relationships by CLay (e.g. 1957) and TIMMERMANN (963), the wing molt by STRESEMANN (1963) and the mor- phology of avian sperm by MGFARLANE (1963). Mayr (1959) has also reviewed this history and has cited additional examples. Each new study has been presented with the hope, and sometimes with the conviction, that here at long last was the key to the classification of birds. But the optimism at the beginning of each new investigation gradually changes into doubt as further data accumulate, Every character even- tually is found to be adaptive, to be subject to convergence and to present inconsistencies in comparison with other cha- racters. The difficulties confronting avian systematists were graphically portrayed in 1959 by STRESEMANN who concluded that the efforts of the past 200 years had failed to provide us with trustworthy information on the relationships of the higher categories of birds. Nevertheless each new set of data has provided us with additional information and it is cer- tainly true that our present understanding of avian rela- tionships is much better than it was 200 years ago. It is also true, as STRESEMANN emphasized, that the relationships of the higher categories of birds are still quite uncertain. So obscure is their phylogenetic history and so uncertain is our knowledge of their genetic relatedness that the orders of birds might almost as well be considered the results of sepa- rate acts of special creation as the products of a monophyle- tic ancestry. The fossil record is unlikely to fill in more than a few of the many gaps for avian fossils are not readily pre- served and hence will never be numerous. In spite of STRESEMANN’S justified pessimism the search continues. Bock (1960) showed that the palatine process of the premaxilla « has little value in showing relationships between families of passerine birds » and thereby helped to free us from certain misconceptions. STRESEMANN, apparently unconvinced by his own argument, continues to produce per- ceptive studies of taxonomic value on wing molt patterns (1963). Although we have béen repeatedly disillusioned we remain Source : MNHN. Paris 114 L/OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE convinced that the evidence exists if only we can learn how to expose it and to interpret it. The search is for new data to supplement the old and our loss of innocent faith has been accompanied by an increasing sophislication in both theory and techniques. There is, in systematics, only one basic technique, that of comparison. Because comparisons between whole organisms present insuperable difficulties it is customary, in fact neces- sary, to compare characters. For characters to be comparable they must be the products of homologous genes. This requi- rement restricts us to comparisons between closely related organisms if we are to avoid the hazards of convergent evo- lution because most aspects of gross morphology are the pro- ducts of many genes some of which may not be homologous in different organisms. To circumvent these difficulties sys- tematists have constantly searched for « conservative » cha- racters which would preserve evidence of evolutionary his- tory in spite of adaptive changes and convergent similarilies. Within the past twenty years the development of « molecular biology » has opened up new possibilities for the comparison of morphological characters clearly determined by homolo- gous genes. Several new techniques for measuring and compa- ring morphological characters at the molecular level of struc- ture have become routine and are available for application to systematies. More complicated techniques which promise much for the future but are not yet ready for general use are being developed. In previous papers (SrBcey, 1960 ; 1962 ; 1964) I have pre- sented the rationale behind the comparative study of homo- logous proteins as a source of data for classification. It may be assumed that all biologists today are aware that the linear sequence of amino acids in a protein chain is a translation of a corresponding sequence of genetic coding units in a seg- ment of DNA. Thus a protein molecule is the direct pheno- typie expression of a gene. The fact that the genetic code has, so far, proved to be universal provides us for the first time with a common unit of genetic curreney, the coding unit of DNA. Because the sequence of genetic coding units is expres- sed as the sequence of amino acids at the protein level it is clear that comparisons among homologous proteins are essen- tially the same as comparisons of the genes themselves. It is also apparent that the ideal basis for comparisons would Source : MNHN. Paris MOLECULAR SYSTEMATICS 115 be the complete amino acid sequences of homologous pro- teins. Unfortunately, the determination of complete sequences is still extremely tedious and is not yet practical for com- parisons among hundreds of species. However, there are seve- ral techniques which can be used in comparative studies of the properties of proleins. Electrophoresis is one such technique which has been used exlensively to characterize such protein systems as mammalian blood serum (JonNson and Wicks, 1959), avian egg white (Sicey, 1960), the plasma proteins of amphibians and repliles (DESSAUER and Fox, 1956) and the muscle proteins and hemoglobins of fishes (Tsuyuki, RoBErTs and VANSTONE, 1965). Electrophoresis is defined as « the movement of charged particles suspended in a liquid under the influence of an applied electric field ». The speed of migration of a protein molecule depends primarily upon its net electrical charge, perhaps to a slight extent upon the distribution of the charged amino acids and, in such media as starch gel, upon the size and possibly the shape of the molecule. The various proteins in a mixture like egg while move at different speeds and thus they separate during elec- trophoretic analysis. By carrying out the separation in a sup- porting medium such as filter paper or starch gel it is pos- sible to produce à dyed pattern indicating the positions of the different proteins after a standard period of time. The pat- terns may then be compared and differences and similarities among species noted and assessed. Electrophoresis is a useful and powerful tool but its limi- tations should never be forgotten. Two proteins with different amino acid sequences can show identical electrophoretic beha- vior and two proteins differing by but a single amino acid may show quite different behavior. One safeguard against erroneous interpretations due to such problems is to use elec- trophoretic comparisons only for protein systems such as blood serum and egg white which contain an array of many different proteins. The complex patterns produced are likely to be completely identical only if they are derived from gene- tically very similar organisms. Another safeguard is to make separate comparisons using different buffer systems since the electrophoretic behavior of a protein varies with the pH of the environment, During the past ten years we have used electrophoretic techniques to compare the egg white proteins of nearly Source : MNHN. Paris 116 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE 2000 species of birds. We have studied the egg white proteins of all of the 27 orders and of 146 of the 170 living families recognized by WErmMoRE (1960). In most cases the protein data support the present classification. The Anatidae and the Galliformes prove to be closely-knit groups as was already clear from the evidence of morphology and hybridization. Similarly the Charadriiformes prove to be a natural unit and the New World nine-primaried oscines also prove to be as similar in their proteins as in their gross morphology. The departures from strict agreement also provide some interesting aspects. The egg white proteins suggest that Stru- thio and the Casuariiformes are related and that Rhea may also be part of the same ancient but monophyletic assemblage. Gavia may be closer to the Laridae than to any other living group and the Alcidae are clearly allied to the other Charadriiformes, not to the penguins as was suggested by VERHEYEN (1958). The Pelecaniformes are remarkably hetero- geneous and the protein evidence raises a strong suspicion that this group may be polyphyletic. The Ciconiiformes howe- ver do show a constant pattern even though there is conside- rable variation. The flamingos are apparently closer to the herons than to the ducks. The Coraciiformes are also probably polyphyletie for the rollers (e.g. Eurystomus) are remarkably unlike Merops, Momotus, Todus and Megaceryle. Among the passerines the Alaudidae, Hirundinidae, Corvi- dae and most of the nine-primaried oscines stand out as uni- form, natural units. The egg white patterns of the Parulidae, Icteridae, Thraupidae (including Tersina) and Emberizinae are essentially indistinguishable. The Carduelidae are close to this assemblage and the Drepanininae may well be considered to be carduelid. Fringilla itself differs sufficiently to present a question about its affinities as has been recognized by Mayr, ANDREW and Hinpe (1956). The Ploceidae and Estrildidae are easily separable from one another and the latter are dis- tinct from the Carduelidae. Complete data supporting these general remarks, and fur- ther data and discussion will be found in Sr8cey (1960 ; in prep.) and SiBLEY and AHLQUIST (in prep.). We have also studied the electrophoretic behavior of the hemoglobin of about 800 species representing all but three Source : MNHN. Paris MOLECULAR SYSTEMATICS 117 of the 27 orders of WEermorE (1960). Most groups of birds have two hemoglobins, a few have one, several have three and a few have four. The oscines are impressively uniform, all have two hemoglobins and the evidence obtained by exami- nation of their behavior under different conditions of pH indi- cates a high degree of similarity throughout. The sub-oscines also have two hemoglobins which are somewhat more hetero- geneous in mobility. The hemoglobins of the non-passerines tend, in general, to corroborate the evidence obtained from the egg white pro- teins. The ostrich and emu have similar hemoglobins and Rhea, although showing some differences, is similar to them. The Podicipediformes, Sphenisciformes, Procellariiformes, Charadriiformes and Gruiformes all have three hemoglobins and, in general, they also show similarities in their egg white patterns. This suggests an ancient relationship which is already generally assumed by systematists. The birds of prey provide some fascinating problems. The Cathartidae have four hemoglobins, the Accipitridae, Pandion and Sagittarius show three and the Falconidae two. It is espe- cially intriguing to find that Falco is more like Tyto than it is like any of the other diurnal birds of prey ! This of course would agree with the suggestions of a falcon-owl relationship made by Srirck and Barnixor (1954), Srärcr (1959) and Vorrio (1955). In his classification BERLIOZ (1950) placed the Strigiformes and Falconiformes close to one another. These are but a few of the results to date from the hemo- globin studies. Complete data and discussion may be found in SIBLEY and BrusH (in prep.). In addition to the studies on egg white and hemo- globin we have also completed an analysis of the eye lens proteins of several hundred species using starch gel electro- phoresis. RABAEY (1959) and Gysezs (1964; 1965) have published extensively on their studies of lens proteins using agar gel electrophoresis. Our results will be presented elsewhere (SIBLEY and BRUSH, in prep.). Most of the taxonomic studies to date have been based upon intact proteins or protein systems such as egg white, blood serum, eye lens proteins or hemoglobin. However, because the sequence of amino acids in a protein is the expres- sion of a segment of the genetic code it is clear that a Source : MNHN. Paris 118 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE technique that will provide an index Lo that sequence should be a useful method for systematies. The simplest approach to the next level of structural complexity is Lo break the protein chain at specific points to produce a set of homologous frag- ments or « peptides ». For example, the enzyme trypsin will split a protein chain only at the positions oceupied by the amino acids arginine and lysine. (This is analogous to divi- ding a sentence into phrases by cutting it at each € a » and each « 1 ».) The resulting peptides may then be characterized by one of several techniques and the behavior of homologous peptides can be compared. One method is to separate the peptides by ion-exchange chromatography and to compare the complex curves which can be produced using automatic ins- trumentation. 1 have reported on one such study which demonstrated that even closely related species possess diffe- rences in the structure of such proteins as ovalbumin and ovo- mucoid (SrBLey, 1964). A study of the tryptic peptides of the ovalbumins of several species of pigeons (Columbidae) has been completed by Corn (1965) using two-dimensional thin- layer chromatography and electrophoresis. This technique has been described by RrrscHarD (1964). In the near future we expect to begin à comparative study of the tryptic peptides of avian hemoglobins. At the next level of complexity, that of peptide composi- tion, the amount of work involved increases sharply but the information is also greatly amplified. Hizz and BUETTNER- Januscn (1965) have demonstrated the value of such studies in their interesting work on the hemoglobins of primates. Further improvements in automatic instrumentation will eventually speed up the processes involved but until they do it will be necessary to restrict this technique to studies invol- ving few species. To determine the complete sequence of the amino acids in even a small protein of less than 100 residues is a slow and difficult process. Several different peptide cleavages must be made and compositional analyses must be carried out on all the peptides to provide the data required for the determina- tion of the sequence. So far the only taxonomically significant studies at this level are those of Smrrx and MARGOLIASH (1964) on eylochrome c and these involve only eleven species and relate to very general problems. Ever since it became certain that genetic information was Source : MNHN. Paris MOLECULAR SYSTEMATICS 119 encoded in the linear sequence of the nucléotides in DNA it has been obvious that an index to that sequence would be the ultimate systematic technique. We are still a long way from being able {o compare the actual nucleotide sequences of homologous DNA segments but the development of the tech- nique of « DNA hybridization » promises to provide an index to the genetic relatedness of any two species. This technique, which has been made applicable to the systematies of higher organisms by the agar-embedding method of Hoyer, McCar- THY and BoLron (1964), provides an index to the base pair complementarity of the two DNA's to be compared. It is thus an index to the degree of similarity between the total genomes of the two species being compared. DNA hybridization promises much for the future but it is a relatively difficull procedure which requires specialized instruments. Avian DNA is easily obtained from the nuclei of the red blood cells and we now have à collection of purified DNA specimens from over 500 species representing all but three of Wetmore’s 27 orders. This material will be utilized in systematie studies over the next few years. What then, in general, has been accomplished in the past 10 years in this field and what may we expect in the future ? Old techniques have been refined and new ones have appeared frequently. Better instruments are constantly becoming avai- lable and yet much better ones are still needed. Collections of proteins and DNA, represenling many groups of organisms, have been accumulated, although many more specimens, espe- cially or fresh material, are needed. So far the results, in Lerms of new and more accurate classifications, are not impressive. This is due partly to the limited collections available and partly to the fact that most of the available techniques are actually relalively crude. However these problems will be overcome as collections grow and techniques improve. The basic validity of the theoretical framework is unchallenged and seems well-founded. This means that the important discoveries and contributions of the molecular approach to systematies still lie in the future and hence an attempt to look ahead is justified. It is reasonable Lo expect that the number of studies using molecular techniques will increase and additional groups of organisms will be investigated. Automatic instruments of great sophistication will be developed for use in molecular Source : MNHN. Paris 120 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE biology and will be applied to comparative studies. They will be costly and they will require skilled technicians to operate them but they will provide vast amounts of new data. The collection and preservation of specimens of proteins and DNA must be greatly improved and expanded. Special expeditions equipped to preserve material in the deep cold of liquid nitrogen will be required. Molecular systematists must be trained in classical taxo- nomy — they must in fact be classical taxonomists — and they must be familiar with the organisms they are studying as living plants and animals. In addition they should be at least reasonably familiar with the concepts and techniques of molecular biology and biochemistry. Finally, let it be clearly understood that the application of the methods of molecular biology to systematics does not insure the solution of all of our problems. The new tech- niques provide new kinds of calipers which can measure pre- viously unavailable characters but the interpretation of the data still requires a systematist who knows, appreciates and understands the other available information about the group of organisms he is studying. The molecular data are enor- mously exciting, and hold great promise for future disco- veries, but they must be viewed as additions to, not substi- tutes for, what is already known about the genetic relation- ships and evolutionary history of plants and animals. ACKNOWLEDGMENTS The work upon which this paper is based was carried out at Cornell University and supported by grants from the Natio- nal Science Foundation (G-4562, G-5859, G-18958, GB-2229) and the U.S. Public Health Service (GM-6889). Acknow- ledgments for technical assistance and specimens will be found in the complete reports cited in the text. RESUME Leur structure étant liée à celle de l'ADN, les protéines peuvent être considérées comme l'expression phénotypique des gènes. Leur comparaison s’avère donc susceptible d’ap- Source : MNHN. Paris MOLECULAR SYSTEMATICS 121 porter des éléments valables du point de vue systématique. Ainsi s’est développée l'étude des protéines par électropho- rèse (étude des déplacements sous l’action d’un champ élec- trique de particules électriquement chargées en suspension dans un liquide). Pour minimiser les sources d'erreur, outre que l’on renouvelle les expériences à pH différent, on utilise des matériaux tels que le sérum du sang ou l’albumen de l'œuf qui contiennent un assortiment varié de protéines. Les schémas de la répartition de ces dernières à l’électrophorèse ne seront identiques que si elles dérivent d'organismes géné- tiquement très proches. Un résumé est donné des résultats obtenus par l’auteur et d’autres chercheurs à la suite d’études des protéines de l’albumen de l'œuf, de l’hémoglobine et du cristallin. Afin de serrer davantage le problème des relations entre espèces, on peut analyser par un procédé chromatographique les divers peptides obtenus par fractionnement des chaînes protéiques à l’aide d’enzymes appropriés permettant de con- naître à quels niveaux se produisent les ruptures. 11 apparaît même possible d'envisager des études taxono- miques fondées sur la séquence complète des amino-acides composant les protéines. Plus encore, des comparaisons des ADN vont être entreprises, le matériel étant actuellement recueilli. Indéniablement, les résultats obtenus par la biologie molé- culaire et la biochimie viennent utilement compléter, sans prétendre les remplacer, les informations recueillies jusqu’à présent par la systématique classique. BIBLIOGRAPHY Amss, P. 1965. — The morphology of the syrinx in passerine birds : its application to the taxonomy of the order Passeriformes. Thesis, Yale University. Bgecuer, W. J., 1953. — A phylogeny of the oscines. Auk, 70 : 270-333. 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G, 1964 — The characteristies of specific peptides from single proteins as data for classification : pp. 435-450 in l'axo- nomie Biochemistry and Serology, ed. by C. A. Leonr, Ronald Press. Six, C. G. (ln prep). — A taxonomic study of the electrophoretie patterns of the egg white proteins of passerine birds, Sex, C. G, and J. E. AniQuisr (In prep). — A taxonomic study of the electrophoretie patterns of the egg white proteins of non- passerine birds. Sues, C. G,, and A. H, Bnusu (In prep). — An electrophoretic study of avian hemoglobins. Smuer, C. G, and A. H. Bnusn (in prep). — An electrophorctie study of avian eye lens proteins. Suirm, E, L, and E. ManGoztasm, 1964, — Evolution of cytochrome c. Fed. Proc., 23 : 1243-1247, Srauucur, W. B, 1954, — Myology and serology of the avian family Fringillidae, Publ. Mus. Nat, Hist. Univ. Kansas, 8(2) :157-211, 23 figs. 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B. 1954. — A systematic study of the avian family Frin- gillidae based on the structure of the skull. Misc. Publ. Mus. Zoo. Univ. Mich., no. 81 : 63 pp. Source : MNHN. Paris 124 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE reuvu, H, E. Rosenrs and W. E. Vansrowr, 1965, — Comparative Sone electropherograms of musele myogens and blood hemoglo- ins of marine and freshwater vertebrates and their application to biochemical systematics. J. Fish. Res. Bd. Canada, 22 : 203-213. Wermons, A, 1960. — A classification for the birds of the world. Smithson. Mise. Coll, 139, no. 11. Venneven, R, 1958. — Analyse du potentiel morphologique et project d'une nouvelle classification des Charadriformes. Bull. Inst. Roy. Sci, Nat. Belgs, 34(18). Vorrro, P., 1955. — Muutuva lintujen järjestelma: System der Vôgel). Ornis Fennica, 32 :108- (Das veränderliche 29. Source : MNHN. Paris DIE MAUSER DES KAMPFLAÂUFERS PHILOMACHUS PUGNAX von Erwin STRESEMANN Praesident der Deutschen Ornithologen-Gesellschaft Président du VIII Congrès International d'Ornithologie (Oxford 1934) und Vesta STRESEMANN Die Geschlechter unterscheiden sich beim Kampfläufer in vieler Hinsicht weit mehr als bei allen anderen Limicolen. Jedermann weiss, dass das Männchen viel grôsser ist als das Weïbchen, dass nur das Männchen ein sehr auffälliges « Hoch- zeitskleid » trägt und ferner, dass es sich nach der Copula weder um die von ihm befruchteten Weïbchen noch um die Nachkommenschaft kümmert. Unbekannt aber war bis vor kurzem, dass sich die Geschlechtsunterschiede auch auf den Zeitpunkt und den Ver- lauf des Gefiederwechsels erstrecken. Erst Frau Elisabeth KozLowa (1956, 1962) hat die Mauser von Philomachus gründlich untersucht und dabei entdeckt, dass dieser Vogel den grôssten Teil seiner Kôrperfedern zwischen zwei Brutcyclen nicht wie bis dahin angenommen worden war zweimal, sondern dreimal erneuert. Das hat sie gefolgert aus dem sorgfältigen Studium vieler Bälge des Zoo- logischen Instituts zu Leningrad, die im Raum der Sowjet- Union in den Monaten März bis Anfang Oktober gesammelt worden waren. Was für Veränderungen das Gefieder im Win- terquartier durchmacht, konnte Frau Kozlowa nur vermuten. Kollegiale Hilfsbereitschaft bat es uns ermôglicht, diese Lücke zu schliessen und von der Mauser des Kampfläufers eine deutlichere Vorstellung zu gewinnen. Wir untersuchten nahezu 90 im Zustand der Mauser gesammelte Bälge, die sich auf die Monate Juli bis Mai verteilen. Darunter sind gegen 50 Exemplare aus dem afrikanischen Winterquartier ; sie wurden uns für unseren Zweck von den Verwaltern der Museen in Bulawayo und Durban freundlichst züugesandt. L'Oiseau et R.F.O. V. 85, 1965, n° spécial. Source : MNHN. Paris 126 L’OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE Es ist uns eine grosse Freude, die wesentlichsten Ergebnisse unserer Philomachus-Studien unserem langjäh- rigen Freund Jaques Berlioz bekanntgeben zu kônnen, bevor unsere Monographie der Vogelmauser abgeschlossen ist, worin das Thema ausführlich behandelt werden wird. Das Brutgebiet des Kampfläufers nimmt den ganzen Nor- den des eurasiatischen Festlandes ein und reïicht bis in die arktische Tundra. Die meisten Populationen, zum Teil auch die ostsibirischen, überwintern in Afrika, wo sich besonders viele Kampfläufer von September bis März südlich des Aequa- tors, bis hinab zur Kap-Provinz, aufhalten. Im zeitigen Früh- jabr wandern sie wieder zu den Brutarealen. Die Vortrupps der Heimkehrer bestehen vorwiegend aus Männchen. Sie pile- gen, bald von der Hauptmasse beider Geschlechter gefolgt, in Holland in der zweiten Märzhälfte, in Ostpreussen gegen Mitte April, in der arktischen Tundra aber erst gegen Ende Mai oder noch später zu erscheinen. Am Südrand des Brutareals werden die Eier im Mai oder in der ersten Juni-Hälfte, in der Arktis von Mitte Juni an gelegt. Die adulten Männchen verlassen das engere Nistareal in Deutschland spätestens in der zweiten Junihälfte, die Weib- chen verbleiben dort, bis ihre Jungen fliegen kônnen. MAUSER DER ADULTEN MANNCHEN Sehr bald nachdem die Männchen aufgehôrt haben die Turnierplätze aufzusuchen (in Deutschland gegen Mitte Juni) streichen sie gesellig umher. Sie beginnen dabeïi ihren Kopf- schmuck und die langen Kragenfedern zu verlieren und auch am übrigen Kôrper das Brutkleid gegen ein schlichtes Rei- sekleid (das postnuptiale Schlichtkleid) zu vertauschen. Aber diese der Wanderung nach Afrika vorausgehende Mauser érstreckt sich nicht nur auf alle Kôrperfedern, sondern auch auf alle Schwung- und Schwanzfedern oder wenigstens die meisten davon. Ein von uns untersuchtes Männchen aus Hol- land hat schon am 17. August Flügel und Schwanz vôllig erneuert. Andere Männchen unterbrechen die begonnene Flü- gelmauser, wenn sie in < Zugstimmung » geraten sind, und beenden sie erst in Afrika. In der afrikanisehen Winterherberge wird, wahrscheinlich zwischen November und Februar, das Kôrpergefieder aber- Source : MNHN. Paris DIE MAUSER DES PHILOMACHUS PUGNAX 127 mals erneuert. Danach trägt der Vogel das zweite Schlicht- kleid (— praenuptiale Kleid). Es kann dem ersten Schlicht- kleid entweder gleichen oder etwas kontrastreicher gefärbt sein. Mit seinem neuen Reisekleid angetan wandert das adulte Männchen im März oder April zur Palaearktis. Erst wenn der Vogel das Ziel seiner Wanderung erreicht hat oder ihm nicht mehr fern ist, verliert er in kurzer Zeit alle Kôrperfedern (vielleicht ausser denen der hinteren Bauchregion), an deren Stelle die ganz anders gefärbten und an Kopf und Hals auch ganz anders gestalteten Federn des Brutkleides hervorwach- sen. Die meiïsten inneren Armschwingen (die Schirmfedern) und zwei bis drei mittlere Steuerfeder-Paare kônnen schon vor Beginn des Frühjahrszuges in Brutkleid-Färbung fertig- gestellt sein ; die etwa verbliebenen Schirmfedern werden auf der Wanderung erneuert (die Schirmfedern mausern dann zum dritten Mal, die Steuerfedern zum zweiten Mal). Das männliche Brutkleid wird also nur zwei Monate lang (in Mit- tel- und Westeuropa etwa von Mitte April bis Mitte Juni) getragen. MAUSER DER ADULTEN WEIBCHEN Ganz anders als bei den Männchen verläuft die Mauser bei den Weïbchen. Sie beginnen frühestens im Juli, meistens erst im August, das Brutkleid mit dem ersten Schlichtkleid (Postnuptial- Kleid) zu vertauschen, mausern dabei aber nicht das Grossge- fieder, sondern wandern Ende August oder im September mit dem alten Flügel und Schwanz nach Afrika. Dort fangen sie frühestens Ende September, in der Regel aber erst im Okto- ber an, Schwingen und Schwanz zu erneuern, und sind Ende Januar damit fertig. Von Dezember bis Anfang Februar wird das Kôrpergefieder mitsamt den Schirmfedern abermals erneuert. Aus dieser heftigen Mauser geht ein zweites Schlichtkleid (das Praenuptial-Kleid) hervor, das sich vom ersten wenig oder gar nicht unterscheidet. Milte oder Ende März, also kurz vor dem Aufbruch aus Afrika zur Palaearktis, setzt die dritte Kôrpermauser ein ; es wachsen nun die kon- trastreichen Federn des Brutkleides. In diese Mauser werden einbezogen das mittlere und in der Regel auch das zweite 9 Source : MNHN. Paris 128 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Wenn das adulte Weïbchen am Nistort eintrifft, trägt es also in der Regel schon das fertige Brutkleid. MAUSER DER JUNGEN oder sogar das dritte Paar der Steuerfedern und wiederum die Schirmfedern. Die Jungen befinden sich noch im vollständigen Jugend- kleid, wenn ihr Zugtrieb erwacht. Sie beginnen das Kôrper- gefieder fast stets erst während der Wanderung oder kurz nach der Ankunft im Winterquartier zu erneuern. Spätestens Ende November tragen sie in der Regel schon ein vollstän- diges Schlichtkleïd. Dieses wird von Ende Dezember ab ganz allmählich in ein ebenso gefärbles zweites Schlichtkleid gemausert, wobei auch die fünf Schirmfedern zum zweiten Mal erneuert werden. Unvermausert bleiben im ersten Win- ter die Handschwingen, die zehn äusseren Armschwingen und alle Steuerfedern. Wenn die Jungen im Frühling ihres zweiten Kalender- jahres wieder in der Heimal erscheinen, tragen sie also noch immer ihren ersten Flügel und in der Regel auch noch das zweite Schlichtkleid. Manche jungen Weiïbchen beginnen während der Wanderung einen Teil des Kôrpergefeders, eine oder zwei Schirmfedern und das mittlere oder die beiden mittleren Paare der Steuerfedern abzustossen und durch typische Brutkleidfedern zu ersetzen. Wie es sich mit der Kürpermauser junger Männchen verhält, wenn sie das Hei- matgebiet erreicht haben, vermochten wir nicht genau fest- zustellen. Wir vermuten, dass sich viele junge Weibchen in ihrem zweiten Kalenderjahr noch nicht fortpflanzen, und dass die gleichaltrigen Männchen zwar auf den Turnierplätzen erschei- nen, aber nur als Zuschauer und ohne dort zu kopulieren. Trscucer (1941) hat zu dieser Frage bemerkt : « Nachdem der Frühjahrszug in den letzten Mai-oder Junitagen beendet ist, trifft man auch im Juni fast stets einzelne Kampfläufer am Kinkeïmer See (Ostpreussen) an, meist Weibchen, biswei- len sogar in ganzen Flügen. Wahrscheinlich handelt es sich meist um einjährige Vôgel. Ein Weïbchen vom 8. Juni 1908 hatle ganz unentwickelten Eierstock. » Die jungen Männchen mausern ihren ersten Flügel im weiteren Brutgebiet, nachdem sie ihn ein Jahr lang getragen Source : MNHN. Paris DIE MAUSER DES PHILOMACHUS PUGNAX 129 haben, beginnen damit also zur gleichen Zeit wie die adulten Männchen, nämlich im Juli. Die jungen Weibchen müssten, um die Schwungfedern gleichzeitig mit den adulten Weïbchen zu erneuern, die weite Entfernung zwischen der nôrdlichen Palaearktis und dem afrikanischen Winterquartier zum drit- ten Mal mit ihrem ersten Flügel bewältigen. Das tun sie nicht, sondern sie fangen schon vor der Wanderung, im Juli, mit der Mauser von Flügeln und Schwanz an, die beim Erwachen des Zugtriebes zum grôssten Teil, wohl zuweilen sogar ganz beendet ist. Auch das Kôrpergefieder wird schon zu dieser Zeit erneuert. RESUME La mue des Chevaliers combattants adultes diffère, pour les deux sexes, de celle des autres Limicoles par le fait qu’il y en a trois et non une ou deux dans l’année. La séquence est la suivante : Plumage nuptial, 1* Plumage neutre (Plumage postnuptial), 2° Plumage neutre (Plumage prénuptial), Plu- mage nuptial. Le mâle adulte mue les rémiges et les rectrices avant de partir dans ses quartiers d'hiver, contrairement à ce que fait la femelle adulte. Il ne reprend la livrée nuptiale que lorsqu'il se trouve dans son aire de reproduction ou qu’il en est très rapproché dans sa migration printanière. Au contraire, la femelle adulte revêt le plumage nuptial peu avant la migration prénuptiale alors qu’elle est encore en zone d’hivernage. Les jeunes des deux sexes reviennent en zone de repro- duction au printemps de leur deuxième année et, en même temps que les mâles adultes, y muent les ailes dès juillet, après avoir porté ces plumes pendant un an. Vraisemblable- ment se reproduisent-ils pour la première fois lorsqu'ils sont dans leur troisième année. LITERATUR Kozzowa, E: (1956). — In : Zool. Journal, 35, pp. 1908-1910 (russ.). Kozrowa, E. (1962). — Regenpfeifervôgel (Charadriiformes), Unter- ordnung Limicolae, in : Fauna der U.S.S.R., Band II, Heft 1, Teil 3. Moskau und Leningrad (russ.) TiscmLer, F. (1941). — Die Vôgel Ostpreussens. Kônigsberg und Berlin. Source : MNHN. Paris THE TRANSEQUATORIAL MIGRATION OF THE MANX SHEARWATER (PUFFIN DES ANGLAIS) by A. LANDSBOROUGH THOMSON Président du XI Congrès International d'Ornithologie (Bâle 195%) The main purpose of this paper is to draw attention to some theorelical implications of the transequatorial migra- tion of the nominale subspecies of the Manx Shearwater Puf- finus puffinus (Brünnich). Important new information about this movement has been acquired in the last dozen years as a result of continued large-scale ringing in the British Isles. These recent data are here presented as a whole, having hitherto been published only in scattered form (THOMSON & LeAcn 1952, 1953 ; SPENCER 1955 et seq.) or so far not at all. The records showing transequatorial movement naturallÿ constitute only a small fraction (2.6 per cent) of the total ; 115 712 birds of this species had been ringed under the Bri- tish national scheme up to the end of 1963, and had yielded 1480 recovery records in all. Less than half of these records, including only two transequatorial recoveries, were available at the time of an analysis, embracing observational data as well, which showed that many of the birds breeding in the southern part of the Irish Sea make feeding visits to the Bay of Biscay during the period March-August (Lockrex 1953). A further analysis of limited scope has recently been made (Harris in press, in litt.). The present writer is indebted to W. R. P. Bourne (in litt.) for some valuable suggestions incorporated here. The nesting colonies, mainly insular, of Puffinus puffinus puffinus are distributed from Iceland and the Faeroe Islands, through the Bristish Isles and north-western France (Finis- tère), to the Atlantic islands from Madeira to the Azores (formerly also Bermuda). In the British area, there is a major breeding concentration on islands in the southern part of the Irish Sea, notably Skokholm (off Pembrokeshire, Wales) ; the species no longer inhabits the Isle of Man, from which it L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris MIGRATION OF THE MANX SHEARWATER 131 takes its English name, but there are nesting colonies off the coasts of Ireland and northern Scotland. There are too few records to show whether the more northerly of these British populations behave differently from those in the Irish Sea. RINGING DATA In the following table, the records in each category are given in chronological order of recovery dates in successive years of life since that of ringing (irrespective of calendar years). Where an explicit recovery date is not available, the date of report is given in brackets. TRANSEQUATORIAL AND TRANSOCEANIC RECOVERIES OF MANX SHEARWATERS RINGED IN THE BRITISH ISLES All ringed on the island of Skokholm, off Pembrokeshire, Wales (51°42'N., 5°16"W), except as otherwise stated : (B) = ringed on Bardsey Island, Caernarvonshire, Wales ; 52°45°N. 4°48°W. (C) = ringed on Copeland Island, co. Down, Northern Ireland ; 54°40°N. 5°31°W. (Sr) — ringed on Skomer, Pembrokeshire, Wales ; 51°43%N., 5°16W. Ringed as nestlings (pulli) RECOVERED IN SOUTHERN HEMISPHERE WITHIN A YEAR Brit. Mus. Date (Wat. Hist) ringed Date and Place of Recovery EC22980 2.9.62 16.10.62 Caraguatatuba (Säo Paulo), Bra- zil; 23°38S., 45°32°W. EC05827 31.8.61 18.10.61 Bay of Guanabara (Rio de Ja- neiro), Brazil ; c.22°50'S., 43°12"W. ECO08187 6.9.61 <e. 20.10.61 Porto Bello (Santa Caterina), Bra- zil; 27°05S., 48°35"W. EC52449 27.8.64 24.10.64 Niteroi (Rio de Janeiro), Brazil; Lente ha EC56899 79.64 24.10.64 (Santa Caterina), Brazil ; 48°39"W. AT37976 7.9.55 25.10.55 Santos (So Paulo), Brazil ; 23°58S., 46°20°W. EC56797 7.9.64 25.10.64 Camboriä (Santa Caterina), Bra- zil; 27°01’S., 48°38"W. Source : MNHN. Paris 132 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Brit, Mus. Date Wat. Hist) ringed Date and Place of Recovery EC42459 7.9.63 29.10.63 ratuba (Parana), Brazil ; OS. 48°36/W. AT37604 29.855 29.10.55 nr. ltanhaem (Säo Paulo), Brazil ; 2427S., 4644W. EC56748 9.9.64 211.64 nr, Laguna (Santa Caterina), Bra- zil; 28°15'S., 48°407W. ECO568— 27.8.61 (81161) Bay of Guanabara (Rio de Ja- neiro), Brazil. 3063556 15.11.58 Caraguatatuba (Sto Paulo), Brazil. AX4904 2011.51 Rio de Janeiro, Brazil; 229558, 4312W. AT37929 22.11.55 Rio de Janeiro, Brazil. AT38100 2411.65 nr. Itanhaem (So Paulo), Brazil. EC23580 25.11.62 Bay of Guanabara (Rio de Ja- neiro), Brazil. EC22110 2511.62 Praia de Lagoinha (Santa Cate- rina), Brazil ; 27°48S., 48°33"W. EG07739 49.61 712.61 Itanhaem (Sio Paulo), Brazil. EC22608 29.62 1212.62 Sto José do Norte (Rio Grande do Sul), Brazil; 3159. 5204 W. AT36220 25.856 (C) 2112.56 Albardo (Rio Grande do Sul), Brazil; 33195, 6242 W. AT24758 23.854 —.? 12 nr. Porto Alegre (Rio Grande do Celate»).54 Sul), Brazil; 30°308, 50°40°W. EC07527 3.9.61 10. 4.62 nr. Macedo (Buenos-Aires), Argen- tina ; 37°16S., 57°05"W. REGOVERED IN SOUTHERN HEMISPHERE IN SUBSEQUENT YEARS EC23890 11.962 29. 9.63 Sio Sebastiño (Sio Paulo), Brazil ; 22°49S., 44°09W. 2057768 9.9.60 e. 22.11.61 nr, Venus Bay, South Australia ; (long dead) 33108, 134°28E. AT25087 5.954 2211.55 Florianopolis (Santa Caterina L), Brazil; 27°32 48°30°W. AT72646 31.8.60 (Sr) (10. 2.65) nr, Aracati (Cearä), Brazil 4°32S., 37°45°W. Re&covERED 1N NORTH-WESTERN ATLANTIC AT50166 16.857 10. 7.58 off Cape Spear, Newfoundland ; AT32N., 52°34,W. ATI1692 26.852 (30. 6.54) Bonavista Bay, Newfoundland ; A8°42,N., 53°107W. Source : MNHN. Paris MIGRATION OF THE MANX SHEARWATER 133 Ringed as adults Brit. Mus. Date (Nat. Hist) ringed Date and Place of Recovery AT48865 19.7.57 22. 9.57 off Cabo Frio (Rio de Janeiro), Brazil; 23018. 42°02"W. SS04518 104.64 (B) 211.64 Santos (Sio Paulo), Brazil. AT22469 1.7.54 12. 4,55 off Santa Barbara (Abrolhos), Bra- zil; 1755, 3839 W. AT16559 2.7.53 1212.54 Itanhaem (Sto Paulo), Brazil. AT28060 15.6.55 2. 2.57) San Francisco do Sul (Santa Cate- rina), Brazil; 26°198., 48°36"W. EC01778 64.61 21.10.62 Praia de Lagoinha (Santa Cate- rina), Brazil. AT16447 1.7.58 2210.55 Itanhaem (Säo Paulo), Brazil. 2051196 30.3.60 e. 24.10.62 Tramandai (Rio Grande do Sul), Brazil; 29°508, 50°03"W. 2052739 284.60 23.12.62 Puerto Coronilla (Rocha), Uru- guay ; 33°507S, 53°307W. 2025165 16.759 18.10.62 nr, Joinville (Santa Caterina), Bra- zil; 26018S., 48°49"W. AT49738 26.757 18.10.62 nr, Joinville (Santa Caterina), Bra- zil. AT8306 19.747 19.10.52 Cablo San Antonio, Argentina ; c. 36°35S., 56°50"W. AT42047 168.56 15.11.62 Itapema (Santa Caterina), Brazil ; 27006S., 48°36"W. ANALYSIS The unique record from South Australia can at present be considered only as exceptional, probably due to the bird being carried downwind from the zone of the westerlies in the southern Atlantic Ocean ; the locality is far outside the otherwise known range of the nominate subspecies, although not very distant from that of the form breeding in New Zealand seas. The two records from Newfoundland relate to the breeding season and presumably to non-breeding birds, but it is mere guesswork whether these had travelled north after « wintering » in South American waters or had perfor- med only a transoceanie movement ; there are also some observational records from off the New England coast. The remaining 38 recovery records, all from South America, may Source : MNHN. Paris 134 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE be treated as a single block of data. They confirm that migra- tion to these waters, earlier rather vaguely known from obser- vational data, is a regular phenomenon. Of these records, 25 are of birds ringed as nesilings ; 21 were recovered in their first year of life, 3 in their second, and one in its fifth. The other 13 are of birds ringed as adults at their breeding stations ; they were then of unknown age, and all that need be said is that the recoveries are spread over seven years from that time but fall mainly in the first three. There was at one stage a bias against longterm recoveries, because the rings corroded too easily ; but the maijority of the birds have been marked with rings of more durable metal. The monthly incidence of the recoveries in South Ameri- can waters is as follows : Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. (late) Ringed as pulli First year Enéouge ‘8782 TETE UE Later years 1 1 1 Ringed as adults FE a A à: + a 2 2 à DE OM PRE CU A ER T The concentration of the records in the period from late September to the end of December is remarkable-34, as against 2 in January-March and 2 in April (second week). The recovery localities are spread along the coasts of Bra- zil, Uruguay, and Argentina between 4° and 38°S. lat. The most northerly record, the only one north of Cape Säo Roque, is a February one and the bird was possibly on pas- sage ; the next is from about thirteen degrees further south. There is then a concentration of 30 recoveries between 20° and 30°S. lat. Of the remaining 6, the most southerly are in 36°35’S. lat. and 37°15S. lat. - the former in October and the latter, strangely, in April. The very first record on the list shows how quickly the birds may cross the equatorial belt ; a nestling ringed on 2 September was already in 23°38'S. lat. by 16 October of the same year. (The first young quit their natal burrows on Skokholm early in September, having been abandoned by their parents towards the end of August.) There are South Source : MNHN. Paris MIGRATION OF THE MANX SHEARWATER 135 American recoveries not only of young birds in the first year of life, but of adults in the year of ringing (and presumed breeding) ; the interval in one such instance was from 19 July to 22 September. There is a noticeable concentration of records in certain calendar years - the northern winters of 1955-56, 1961-62, and 1964-65, but especially that of 1962-63. In the latter part of 1962 there were no less than 11 South American recoveries, of which 5 were of young birds of the year, and 6 of birds ringed as adults more than a year earlier. Furthermore, 5 (including one young bird) of these 11 were recovered (found dead or dying) in the period 16-24 October, 2 of them toge- ther at the same place on the same day and all on the stretch of coast between 23° and 30°S. lat. The information about the circumstances of recovery is in general too meagre to be belpful, but the figures themselves point to local conditions as a cause ; there is certainly no correlation with the statistics of birds ringed in different years. Apart from these positive facts of the ringing data, there is the outstanding negative point that the scarcity of South American recoveries after December is matched by an almost equal scarcity of January and early February recoveries anywhere else. (From observational data it is known that the birds are found in only small numbers in the Bay of Biscay at that time ; and a few of them appear at the breeding sta- tions in the Irish Sea before the end of February.) One might conclude that at the beginning of the year, whether the birds are still in the same South American waters or back in Euro- pean walers, some factor must be militating against the occurrence of recovery records ; but no such factor (behaviour- istic, environmental, or artificial) is apparent. Allernatively, the majority of the birds may be in some different waters where the chances of recovery are much less- possibly fur- ther south in the good feeding area of the upwelling of the Falkland Current, which is not only off an unpopulated coast but in the west-wind zone where bodies would tend to be washed offshore (instead of onshore as further north). DISCUSSION There are of course many examples of extensive transequa- torial migration among both land-birds and sea-birds native Source : MNHN. Paris 136 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE to the higher latitudes of the Northern Hemisphere. The advantages for survival of the species, as against the phy- siological cost and hazards of long journeys, are nevertheless not fully understood, although some instances seem clearer than others. The benefits, as compared with a shorter migra- tion to lower latitudes in the same hemisphere, are presuma- bly one or more of the following : (a) more abundant food of the kind to which the birds are adapted in high northern latitudes ; (b) better climatic conditions for obtaining food and for life generally ; and (c) longer hours of daylight for obtaining food. It is also a commonplace that no similarly extensive jour- neys are made by land-birds native to the Southern Hemi- sphere. This fact is usually considered to reflect the relatively small size and southward extent of the land-masses in the South Temperate Zone, as these provide no real counterpart to the breeding areas available to northern migrants. On the other hand, there are vast southern expanses of ocean, with numerous islands providing breeding places for great colo- nies of sea-birds ; and correspondingly there are several well- known instances of southern sea-birds (all of them in the Pro- cellariiformes) that regularly visit middle or high northern latitudes while it is summer there. Wilson’s Petrel Oceanites oceanicus is an outstanding example (Roserrs 1940 ; et al.) and likewise certain shearwaters Puffinus spp. to be men- tioned more fully below (Wynne-Enwarps 1925 ; KuropA 1957 ; Pazmer et al. 1962 ; Bourne 1963 ; and sources cited for particular species). It has for long been known that the Greater Shearwater Puffinus gravis, breeding in the Tristan da Cunha group in the South Atlantic (c. 37°S. lat.), spends the northern sum- mer in the North Atlantic and reaches as far as the Arctic Circle. The adults desert their young in April, and the young themselves depart in May ; the birds arrive back at the breed- ing stations in late August or in September (ROWAN 1952). Arrival in New England and Newfoundland waters takes place at the end of May or early in June ; one ringed as a nestling has been recovered off Newfoundland as early as 15 June in its first year of life. Many of the birds thereafter spread towards the north-east, and the species has its period of maxi- mum abundance off he west coast of Europe from mid- August Lo mid-October. The dates show that both passages Source : MNHN. Paris MIGRATION OF THE MANX SHEARWATER 137 through the tropics, from which there are indeed few records, must be rapid ; and also that the movement during the sum- mer in the North Atlantic tends to follow a wide loop in a clockwise direction (Voous & VATTEL 1963). The Short-tailed Shearwater P. tenuirostris breeds in the Bass Strait, south-eastern Australia, and migrates to the North Pacific, as shown by observational evidence supported by some ringing results (MarSHALL & SERVENTY 1956 ; SER- VENTY 1953, 1956, 1957 et seq.). There is a strikingly constant annual cycle ; the parents probably migrate in mid-April, the young leaving late in April or early in May, and sexually mature birds return to the breeding stations late in Septem- ber. The pattern of the movement has been described as a figure-of-eight, and the northern part of this is a wide clockwise loop. The main concentration in June-August is in an area extending from southern Kamchatka (48°N. lat.) and the Aleutian Islands through the Bering Strait at least to Wrangel Island (71°N. lat.). Later in the summer and in early autumn there is much movement down the west coast of North America, although it appears that the participants are predominantly immature birds (of which there is a large population, as breeding does not take place until 6 years of age). Passage through the tropics again appears to be rapid, and a bird ringed as a fledgling was recovered off Shikopu, Japan (33°N. lat.), before the end of May in its natal year. The Sooty Shearwater P. griseus has a wider breeding dis- tribution in the Southern Hemisphere than the other two spe- cies just mentioned, and as three main populations are invol- ved the movements are more difficult to interpret. There is nevertheless evidence that migrations performed by many birds of this very abundant species include clockwise loop movements in the northern oceans (BOURNE 1956 ; RICHDALE 1963 ; Purccrrs 1963 ; WarHAM 1964). The suggestion, made by various authors, is that in all these southern species the birds perform a peripheral move- ment round the anticyclones normally present in summer in the centres of northern seas in middle latitudes, being assis- ted by following winds throughout ; such a movement, in the Northern Hemisphere, is naturally in a clockwise direction. One is thus led to speculate whether the migration of the Manx Shearwater may be a counterpart, with a loop migra- tion in the South Atlantic — counter-clockwise, as with anti- Source : MNHN. Paris 138 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE cyclones in the Southern Hemisphere — but of such a route there is no positive evidence whatever, and the species has only once been recorded in southern Africa. There is the fur- ther possibility that the pattern as a whole may be a figure- of-eight, as there seems to be some slight observational evi- dence of a clockwise loop in the North Atlantic, round the anticyclone area of the Azores, in spring. Another northern species of petrel known to make a tran- sequatorial migration of considerable extent is the Storm Petrel Hydrobates pelagicus. The breeding distribution is generally similar to that of the European forms of the Manx Shearwater, but the migration is confined to the eastern side of the Atlantic Ocean and the species does not visit the New World. The chief breeding concentration is probably in the British Isles, but the birds are absent from home waters in winter and also from the northern part of the North Atlantic generally (RaNxIN & Durrey 1948). At that time, on the other hand, they are abundant off the west coast of Africa. It is possible that the majority remain in tropical waters north of the Equator, but many go further south - even to the Cape of Good Hope and beyond. The evidence is almost entirely observational, but there are two African records of birds ringed as adults in the British Isles - one from off Mauretania in the following January (SPENCER 1957b), and the other from Cape Province in the following February (unpublished). Among all birds, so far as is definitely known, the Manx Shearwater is unique in having a regular post-breeding migra- tion from Europe to South America. RESUME Analyse des reprises trans-équatoriales et trans-océaniques de Puffin des Anglais (Puffinus p. puffinus) montrant notam- ment la répartition mensuelle et la distribution géographique de 38 reprises faites au large des côtes de l'Amérique du Sud ; une reprise exceptionnelle dans le Sud de l’Australie et deux autres en été à Terre-Neuve, individus vraisemblable- ment non nicheurs. Il est fait état du faible nombre d’indi- vidus observés en Amérique du Sud, en janvier et février alors Source : MNHN. Paris MIGRATION OF THE MANX SHEARWATER 139 qu’il n'existe toujours aucune certitude de la présence de gros effectifs dans les eaux européennes. Les migrations trans-équatoriales en général et plus par- ticuliérement celles des oiseaux pélagiques sont également commentées. Celle du Puffin des Anglais est comparée à celles de ses congénères du sud ainsi qu’à celles du Pétrel tempête. REFERENCES Bourne, W. R. P., 1956. — Migrations of the Sooty Shearwater. Sea Swallow, 9 : 23-25. Bourne, W. R. P., 1963. — A review of oceanic studies of the biology of seabirds. Proc. XIII Internat. Orn. Congr., Ithaca, N. Y., 1962 : 831-854. Hannis, M. P. (in press). — Breeding biology of the Manx Shearwater Procellaria puffinus. Ibis. Kuropa, N,, 1957. — A brief note on the pelagic migration of the Tubi- nares. Rep. Yamashina's Inst. Orn. Zool., 11 : 436-449. LockLey, R. M. 1953. — The movements of the Manx Shearwater at sea during the breeding season, Brit. Birds, 46 : suppl. 1-48. MansxaLz, A. J., & Servenry, D. L. 1956. — The breeding cycle of the Short-tailed Shearwater, Puffinus tenuirostris (Temminck), in rela- tion to transequatorial migration and its environment. Proc. Zool. Soc. Lond., 127 : 489-510. Pazmen, R. 8. (ed.) et al., 1962. — Handbook of North American Birds, vol. 1. New Haven and London (Contributions by Bourne ; Par- MER ; SERVENTY). Pricuies, J. H., 1963. — The pelagice distribution of the Sooty Shearwater Puffinus griseus. Ibis, 105 : 340-353. RankIN, M. N,, & Durrey, E. A. G., 1948. — A study of the bird life of the North Atlantic. Brit. Birds, 41 : suppl. 1-42. Ricapazx, L. E., 1963, — The biology of the Sooty Shearwater Paffinus griseus. Proc. Zool. Soc. Lond., 141 : 1-117. Roserts, B. B., 1940. — The life cycle of Wilson's Petrel Oceanites oceanicus (Kuhl). Sci. Rep. British Graham Land Exped. 1934- 1937, 1 : 141-194. Rowaw, M. K,, 1952. — The Greater Shearwater Puffinus gravis at its breeding grounds. Ibis, 94 : 97-121. ServenrY, D. L., 1953. — Movements of pelagic sea-birds in the Indo- Pacific region. Proc. 7th Pacific Sci. Congr., 4 : 394-406 (and discussion, 407). Senvenry, D. L, 1956. —A Japanese recovery of an Australian-ringed Puffinus tenuirostris. Ibis, 98 : 316. Senvenrv, D. L, 1957 et seq. — The banding programme on Puffinus tenuirostris. I. First Report. C.S.I.R.O. Wildlife Research, 2: 51-59; and subsequent reports. Source : MNHN. Paris 140 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Srencer, R., 1955, 1957 a and b, 1958, 1959, 1962 a, 1963, 1964. — Report on bird-ringing for 1954; do. 1955, 56, "57, 758, *61, ’62, 63. Brit, Birds, 48 : 461-498 ; and in vols. 50 (two), 51, 52, 55, 56, 57. Srencer, R., 1962 b. — British ringed Manx Shearwater in Australia. Brit. Birds, 55 : 86-87. THowson, A. L., & Leacn, E. P. 1952, 1953. — Report on bird-ringing for 1951; do. 1952. Brit. Birds, 45 : 265-277, 341-357; 46 : 287- 303, 313-330. Voous, K. H., & Warez, J., 1963. — Distribution and migration of the Greater Shearwater. Ardea, 51 : 143-157. Wanna, J., 1964. — Marked Sooty Shearwaters Puffinus griseus in the Northern Hemisphere. — Ibis, 106 : 190-191. Wynne-Enwanps, V. C. 1935. — The habits and distribution of birds on the North Atlantic. Proc. Boston Soc. Nat. Hist., 40 : 233-846. Source : MNHN. Paris DISTRIBUTION REGIONALE ET ALTITUDINALE DES GENRES GARRULAX et BABAX ET NOTES SUR LEUR SYSTEMATIQUE par Charles VAURIE American Museum of Natural History, New York La révision magistrale du genre Garrulax publiée par le Professeur BErLIoZ (1930) a contribué grandement à notre connaissance de la systématique de ce vaste et intéressant groupe des Timaliinés. Elle fut en partie basée sur le grand nombre de spécimens historiques envoyés au Muséum de Paris soit par les missionnaires français dans l’ouest de la Chine, tel que le fameux Père Davip, soit par des explora- teurs français de marque, tel que le Prince Henri d'ORLÉANS. Nombre de formes bien différenciées furent décrites d’après ce matériel par VERREAUX, OUsTALET, ou le Père Davin lui- même. Leurs types sont à Paris. Antérieurement (1928), MEINERTZHAGEN avait publié une utile liste nominale d'ensemble, accompagnée d’une carte, dans son article bien connu sur la zoogéographie des Hima- layas. Il ÿ reconnaissait 42 espèces, 3 de moins que BERLIOZ, mais {ous deux réunirent les genres Babax David, 1875, et Garrulax Lesson, 1831. Depuis la publication de ces deux ouvrages pionniers, des suppléments d'informations furent apportés : par moi-même (959) dans The Birds of the Palearctic Fauna; par RipLey (1961) pour celles des Indes ; et pour l’ensemble par DEIGNAN (1964) dans sa révision pour la « Check-list of birds of the world ». RipLey, DEIGNAN et moi-même reconnaissons Babax, et la distribution des deux genres est illustrée sur la carte (fig. 1). Dans son ensemble, cette carte est semblable à celle de MEINERTZHAGEN, quelques détails mis à part, avec l'exception plus notable qu’elle comprend Ceylan dans l'aire de distribution parce que dans le sud-ouest de cette île habite une espèce endémique (G. cinereïfrons Blyth) que présume- L'Oiseau et R.F.O., V. 35, 1965, n° spécial. Source : MNHN. Paris 142 L'OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE ment MEINERTZHAGEN et aussi BERLIOZ rapportaient aupara- vant à Turdoides Cretzschmar. CONSIDÉRATIONS SYSTÉMATIQUES Cet article est surtout une discussion de la distribution de Babax et Garrulax, et non pas une étude systématique, mais il doit prendre en considération la validité de Babax et les limites spécifiques de certains Garrulaz. Notre compréhension taxonomique de ce groupe d'oiseaux est avancée mais quelques différences d'opinion persistent. Le cas de cinerei- frons vient déjà d’être mentionné. Un autre concerne la posi- tion systématique de phoenicea Gould que Berzioz, MEr- NERTZHAGEN, et aussi Ripcey placent dans le genre Garrulax, alors que DrrGnan place cette espèce dans le genre Liocichla Swinhoe, où elle me semble aussi être mieux à sa place. Cette question avait déjà été étudiée par DELACOUR (1933), qui avait également placé phoenicea dans le genre Liocichla. Babax est clairement apparenté à Garrulax, surtout aux espèces séparées antérieurement sous le genre Janthocincla, mais il est généralement admis que ses trois espèces (lanceo- latus, waddelli, et koslowi) forment un groupe à part; elles se différencient de toutes les autres par leur livrée terne et très lourdement striée. Les deux seules espèces de Garrulax nettement striées (striatus Vigors et lineatus Vigors) ont une pattern des stries totalement différente. Babax diffère aussi de Garrulax par le bec moins turdiforme et beaucoup moins emplumé à la base et sur les narines, et aussi par la forme de la queue plus étagée. C’est dire que les relations systéma- tiques sont à mon avis plus clairement exprimées par l’ad- mission de Babax, quoiqu'il faille admettre que les différences morphologiques mentionnées ne sont peut-être pas très impor- tantes au niveau du genre (voir aussi DELACOUR (1946), qui a reconnu Babar comme formant la transition entre Tur- doides et Garrulax). Les autres espèces, maintenant toutes incluses dans Gar- rulax, étaient antérieurement réparties dans un grand nombre de genres ; DEIGNAN (op. cit.) donne une liste de 18 syno- nymes ! Parmi ceux-ci, Janthocincla Gould, 1835, Trochalop- teron Blyth, 1843, Grammatoptila Reïichenbach, 1850, et Dryo- nastes Sharpe, 1883, ont été les plus usités et certains voudront peut-être les considérer comme sous-genres. Source : MNHN. Paris DISTRIBUTION DES GENRES GARRULAX ET BABAX 143 Il y a aussi des différences d'opinion concernant le nombre des espèces, et il me semble certain que DEIGNAN a réuni trop de formes qui sont véritablement ou probablement des espèces distinctes. Dans le cas des formes paléarctiques que je connais le mieux, il ne me semble pas que lunulatus Ver- reaux, 1871, soit conspécifique de bieti Oustalet, 1897, comme le pense DEIGNAN, car l’état actuel de nos connaissances sug- gère que leurs distributions empiètent l’une sur l’autre dans le Setchouan occidental. Je préfère aussi ne pas suivre Dei- GNAN quand il réunit caerulalus Hodgson, 1836, et poecilo- rhynchus Gould, 1863, parce que leurs livrées sont vraiment trop différentes : tous les caerulatus, quelle que soit leur sous-espèce, ont la gorge et la poitrine d’un blanc pur qui tranche absolument avec les joues et les côtés du cou bruns et les flancs gris et bruns, tandis que toutes les formes de poecilorhynchus ont la gorge et le haut de la poitrine bruns et concolores avec les joues et les côtés du cou, le bas de la poitrine étant gris et se fondant avec les flancs. Les aires de distribution de ces deux oiseaux se rapprochent dans le nord- ouest du Yunnan mais n'ont pas l'air d’empiéter l’une sur l’autre, du moins d’après ce que nous en savons ; mais dans le cas de lunulatus et bieti, qui apparemment se chevauchent, nous devons constater que la coloration du plumage et sa pattern sont similaires, à quelques petits détails près. Ceci devrait nous faire hésiter à réunir des formes dont la colora- tion et la pattern sont très différentes quand il n’y a aucun signe d’intercroisement. Dans le cas de l’Inde tropicale, DEIGNAN considère que nuchalis Godwin-Austen, 1876, et chinensis Scopoli, 1786, sont conspécifiques comme le sont jerdoni Blyth, 1851, et cachinnans Jerdon, 1839, quoique Ripzey, dont j'accepte l’opi- nion, traite les quatre oiseaux comme espèces séparées dans son inventaire des oiseaux de l'Inde. BERLIOZ (1930) avait déjà conclu que les oiseaux des trois paires mentionnées ci-dessus caerulatus-poecilorhynchus, lunulatus-bieti, nuchalis-chinensis), n'étaient pas conspéci- fiques quoique apparentés, mais considérait que jerdoni l'était avec cachinnans. Ces deux derniers ne nichent peut- être pas exactement dans le même district mais habitent des régions très voisines à l’ouest de Madras, ce qui a peut-être influencé Ripcey. Cachinnans a la poitrine rousse et la gorge noire, non striées, mais ces parties sont gris pâle chez jerdoni 10 Source : MNHN. Paris 144 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE et légèrement striées. Ce déplacement de caractère chez deux formes voisines suggère, je crois, qu'elles ne sont pas con- spécifiques. En résumé, j’augmente de quatre la liste des 44 espèces de Garrulax admises par DEIGNAN, soit un total de 48. Il est possible que ce chiffre soit encore augmenté quand nous con- naîtrons mieux la distribution de certaines formes des mon- tagnes de l’ouest de la Chine. Garrulax est donc, de beaucoup, le genre des Timaliinés le plus riche en espèces. DISTRIBUTION RÉGIONALE Le Professeur BERLIOZ (1930) a insisté sur le fait que Garrulax est essentiellement caractéristique de la faune orien- tale, et MEINERTZHAGEN (1928) pense que ce genre a colonisé les Himalayas depuis la Chine occidentale, probablement à partir d'un centre de dispersion situé au Setchouan et au Yunnan. Je partage la même opinion, mais il me semble que le centre de dispersion doive être placé un peu plus au sud, dans le Yunnan et la partie indo-chinoise de la région orien- tale, mais je liens à signaler que Garrulax et Babax, loin d’être restreints à la région orientale, sont aussi très bien représentés dans la région paléarctique. La distribution que nous constatons de nos jours dans cette dernière région dérive peut-être des relictes occasionnées par l'élévation des Hima- layas, ou par une expansion plus récente vers le nord. Sans doute les deux facteurs ont-ils joué et semblent avoir été suivis par un degré bien marqué de spécialion. Dans la région paléarctique, nous trouvons maintenant les 3 espèces de Babax, deux y sont endémiques, ainsi que vingt et une des 48 espèces de Garrulax, dont six sont endémiques. L'une d'elles (lineatus) habite le Turkestan russe, c'est-à-dire bien au nord des puissantes et hautes barrières de l’Indou- Kouch et du Tien-Chan ; elle représente soit une relicte, soit une expansion du nord-ouest des Himalayas. Les vingt autres habitent le nord et l’ouest de la Chine au nord du Yangtsé, c’est-à-dire au nord de la ligne qui sur la carte (fig. 1) indique la limite sud de la région paléarctique. Beaucoup de ces espèces vivent à de très hautes altitudes, et l’une d'elles (davidi) s'étend vers le nord jusqu’à 49° de latitude environ Source : MNHN. Paris DISTRIBUTION DES GENRES GARRULAX ET BABAX 145 dans la Mongolie Intérieure et la Mandchourie de l’ouest ; onze de ces 20 espèces habitent le Kansou et la Tsinghai, et 13 le Tibet ; il est certain qu'aucun de ces pays ne fait partie de la région orientale. J'ai ajouté Babax waddelli aux espèces endémiques de la région paléarctique avec raison quoiqu'il Afghanistan end Rüssion Turkeston Himaloyes China south of 1e Yongtre Asam south ot he Brobmoputro Southern limite cf he Poleatclic Region — = — — — os Number of species per region Fig. 1. — Distribution des genres Garrulax et Babax. Source : MNHN. Paris 146 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE habite aussi l'est des Himalayas, mais seulement aux très hautes altitudes dans l'extrême nord-est du Sikkim, ce qui ne constitue qu'une petite extension du plateau tibétain voi- sin. La liste des espèces qui suit est établie par régions ; elle montre que dix-huit d’entre elles habitent les Himalayas mais qu'une seule (variegatus) est endémique. Toutes les autres habitent aussi la Chine occidentale ou la sous-région Indo- Chinoise, ce qui vient à l'appui de l'opinion avancée par MEI- NERTZHAGEN pour qui Garrulax a colonisé les Himalayas en venant de l’est. La sous-région Indo-Chinoise est la plus riche (32 espèces), ce qui vient à l'appui de ma théorie quand je considère cette région comme le centre le plus actif de spéciation et d’expan- sion. Six espèces habitent l'ile d'Haïnan, et quatre Formose, mais l’une de ces dernières (poecilorhynchus) ne se trouve pas dans la sous-région Indo-Chinoise et provient sans doute de la Chine. A l'exception de l'Afghanistan et du Turkestan, avec leur espèce solitaire (lineatus), toutes les autres régions occupées par Garrulax se trouvent sur la lisière extrême de sa distri- bution et n’ont que peu d’espèces ; la plupart sont endé- miques et paraissent être des relictes. Nous trouvons aussi 6 espèces endémiques dans la sous- région Indo-Chinoise, mais à l'exception de strepitans qui a une distribution assez considérable allant du Laos occidental à la Birmanie de l’est et du sud, les autres (milleti, vassali, yersini, virgatus et austeni) habitent une aire très limitée. Elles sont restreintes à un ou deux petits systèmes monta- gneux en Indochine, Birmanie, ou dans l’Assam, et je crois que nous pouvons les considérer aussi comme des relictes. ESPÈCES PAR RÉGION Mongolie Intérieure et Mandchourie (1) : Garrulax davidi. Kansou et Tsinghai (11) : Babax lanceolatus, B. koslowi, Garrulax davidi, G. sukatschewi G), G. cineracens, G. tunulatus, G. maximus, G. ocellatus, G. canorus, G. sannio, G. elliotii. Tibet (13) : Babax lanceolatus, B. waddelli C), B. koslowi, Garrulax (1) Le nom des espèces endémiques est imprimé en caractères gras. (2) Considéré comme endémique, voir texte. Source : MNHN. Paris DISTRIBUTION DES GENRES GARRULAX ET BABAX 147 albogularis, G. striatus, G. maximus, G. ocellatus, G. sannio, G. linea- tus, G. ellioti, G. henrici, G. affinis, G. erythrocephalus. Chine au nord du Yangtsé, y compris les régions précédentes (23) Babax lanceolatus, B. waddelli, B. koslowi, Garrulax perspicillatus, G. albogularis, G. slriatus, G. maesi @), G. davidi, G. sukatschewi, G. cineraceus, G. lunulatus, G. bieti, G. maximus, G. ocellatus, G. poe- cilorhynchus, G. canorus, G. sannio, G. lineatus, G, elliotiï, G. henrici, G. affinis, G. erythrocephalus, G. formosus. Chine au sud de Yangtsé (21) : Babax lanceolatus, Garrulax perspi- cillatus, G. albogularis, G. leucolophus, G. monileger, G. pectoralis, G. maesi, G. chinensis, G. galbanus, G. cineracens, G. caerulatus, G. poe- cilorhynchus, G. merulinus, G. canorus, G. sannio, G. squamatu G. subunicolor, G. elliotit, G. erythrocephalus, G. formosus, G. milnei Assam au sud du Brahmaputra, Birmanie, Siam et Indochine (32) Babax lanceolatus, G. perspicillatus, G. albogularis, G. leucolophus, G. monileger, G. pectoralis, G. striatus, G. strepitans, G. milleti, G. maesi, G. nuchalis, G. chinensis, G. vassali, G. galbanus, G. deles- serti, G. cineraceus, G. rufogularis, G. ocellatus, G. caerulatus, G. rufi- collis, G. merulinus, G. canorus, G. sannio, G. virgatus, G. austeni, G. squamatus, G. subunicolor, G. affinis, G. ergthrocephalus G. yersini, G. formosus, G, milneï. Himalayas (18) : Babax waddelli, Garrulax albogularis, G. leucolo- phus, G. monileger, G. pectoralis, G. striatus, G. nuchalis, G. deles- serti, G. variegatus, G. rufogularis, G. ocellatus, G. caerulatus, G. rufi- collis, G. lineatus, G. squamatus, G. subunicolor, G. affinis, G. ergthro- cephalus. Afghanistan et Turkestan russe (1) : Garrulax lineatus. Sud-ouest et sud de l'Inde (3) : Garrulax delesserti, G. cachinnans, G. jerdoni. Ceylan (1) : Garrulax cinereïfrons. Formose (4) : Garrulax albogularis, G. poecilorhynchus, G. canorus, G. affinis. Haïnan (6) : Garrulax monileger, G. pectoralis, G. maesi, G. chi- nensis, G. canorus, G. sannio. Malaisie et Grandes Sondes (6) : Garrulax palliatus (Sumatra et Bornéo), G. rufifrons (Java), G. leucolophus (Sumatra), G. lugubris (Malaisie et Bornéo), G. mitratus (Malaisie, Sumatra et Bornéo), G. erythrocephalus (Malaisie), DISTRIBUTION ALTITUDINALE Les oiseaux du genre Garrulax habitent surtout les col- lines ou les régions franchement montagneuses, le plus sou- vent à des élévations moyennes, quoiqu’un groupe d’espèces (G) Dzracour (1946, p. 27) considère que maesi fait part d'une super-espèce avec sérepitans, milleti et ferrarius. Cest dire que fer- rarius Riley, 1930 (qui est considéré comme une sous-espèce de sfre- pitans Blyth, 1855, par Detnan, 1964, p. 357) peut constituer, à mon point de vue, une espèce particulière qui porterait, dans ce eas, le nombre des espèces de Garrulax À 49. Source : MNHN. Paris 148 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE niche à basse altitude, descendant au pied des montagnes et dans les plaines ; seul un autre groupe habite à très haute altitude, montant à la limite des arbres et même au-dessus, dans la zone des buissons arborescents. La variation altitu- dinale de certaines espèces ne semble pas dépasser beaucoup plus de 1.000 mètres, apparemment même beaucoup moins pour certaines, mais pour d'autres (telles que Garrulax cine- G.canorus G.cinereifrens Gnuchalis G.ruficollis G.cineraceus G. rufogularis Grufitrons G. caeruletus G. milneï G. lunulatus &.lanceolatus G. variegatus G.subunicolor G.ocellatus 6. sukotschewi G. bieti 5000 4500m 4000 3500 3000 2500 2000 1500 1000 500 Om ————————_—_—_— on À a mm Rig. 2. — Variation altitudinale dans certaines espèces des genres Babax et Garrulaz Source : MNHN. Paris DISTRIBUTION DES GENRES GARRULAX ET BABAX 149 raceus et G. sannio) elle est énorme, ces deux oiseaux nichant à partir du niveau de la mer sur la côte de la Chine jusqu’à 4.700 mètres dans le nord-ouest du Yunnan. Les trois Babax nichent plus haut que la grande majorité des Garrulax, jusqu’à 4.000 et même 4.850 m : apparemment, pas au-dessous de 3.050 dans le cas de B. waddelli et 3.900 dans celui de B. koslowi, mais B. lanceolatus, qui est beau- coup plus largement distribué que les deux autres, niche aussi en régions tropicales et descend jusqu’à 1.400 mètres. Toutes les espèces des deux genres sont notées comme très sédentaires, ce qui n'exclut pas quelques déplacements locaux limités, et elles peuvent être forcées de descendre beau- coup plus bas pendant l'hiver lorsque les conditions sont inclémentes, Par exemple, dans les Himalayas, la race nomi- nale de G. variegatus niche entre 7.000 et 13.500 pieds d’après RipLey (1961, p. 386), qui nous dit qu’elle a été signalée à 3.500 pieds pendant l'hiver. Dans la liste qui suit, je n'ai donc pas tenu compte de ces records hivernaux en basse alti- tude pour les espèces qui ne sont reconnues nicheuses qu’à hautes élévations. Je mentionne seulement le plus bas et le plus haut record ; sans doute en existe-t-il d’autres un peu plus bas ou un peu plus haut, qui seront fournis par des col- lections futures. Cette liste fut établie d’après des sources diverses et c’est, je crois, la première liste qui donne des records pour toutes les espèces. Les records d'altitude sont assez abondants pour certaines régions montagneuses où l’on a beaucoup collecté, comme dans plusieurs secteurs des Himalayas, mais, comme on peut s’en douter, ils sont moins abondants pour le Tibet et la Chine occidentale, alors qu’ils sont très pauvres pour d’autres régions, à moins même qu’ils ne fassent complète- ment défaut. La littérature sur Garrulax est si volumineuse qu’il m’a fallu me restreindre aux études les plus compréhensives ou les mieux connues. Parmi celles-ci, je mentionne, en plus de mon ouvrage (1959) et de celui de Ripcey (1961), l’étude de MEINERTZHAGEN (1927) sur les Himalayas, la série de LupLow sur les Himalayas de l’est et le sud du Tibet (1927-1928, 1937, 1944, 1950, et 1951), celle de BErLI0Z (1930), les deux par Ricey (1926 et 1931) sur la Chine occidentale, GREENWAY (1933) sur l'extrême nord-ouest du Yunnan, BANGS et PETERS Source : MNHN. Paris 150 L'OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE (1928) sur le Kansou occidental et la Tsinghai orientale, l’in- ventaire des oiseaux de la Chine par CHENG (1958), HACHISUKA et UpaGawa (1951) sur Formose, DEIGNAN (1945) pour le nord du Siam, et les ouvrages généraux sur la Malaisie par ROBINsoN (1928), la Birmanie par SMyTHIES (1953), Bornéo par le même auteur (1960), et sur l'Indochine par DELACOUR et JaBouILLe (1931). J'ai aussi consulté les étiquettes des spécimens dans la collection de l'American Museum of Natural History, et dans certains cas, particulièrement pour les espèces très rares ou très peu connues et pour lesquelles l'information fait défaut, j'ai recherché sur cartes l'altitude des localités où l'oiseau a été trouvé. Les espèces sont classées ci-après en plusieurs catégories qui correspondent à la variation sur le continent quand la même espèce habite aussi une île, comme Formose ou Haïnan, mais ne donne pas de records pour Haïnan. Je n’en ai pas trouvé pour cette île dans le rapport de base par HARTERT (1910) ou sur les étiquettes de la grande série que j'ai exa- miné et qui fut, d’ailleurs, étudiée en grande partie par Har- TERT. À l'exception d'une localité dans la basse plaine et une autre sur la côte où G. chinensis a élé collecté, les seules localités que j'ai pu trouver sur la carte sont la chaîne des « Five Fingers » et le « Mount Wuchi ». Chinensis a été pris aussi dans la chaîne qui a peut-être une altitude moyenne de 1.000 mètres à l'exception du Mont Wuchi qui atteint 1.879 et sur lequel quatre des autres espèces d’Haïnan ont été collectées (G. pectoralis, G. maesi, G. monileger et G. cano- rus), mais pas chinensis. Je n’ai pas d'information sur la sixième espèce (G. sannio) que DEIGNAN (1964) ne mentionne pas pour Haïnan mais qui a été signalée en dehors d'Haïnan par CHENG (1958) et par moi (1959, p. 432) ; malheureusement je n’ai pas gardé de note sur la localité et je n’ai pas pu la retrouver. La variation allitudinale de deux espèces, ou plus, de chaque catégorie est illustrée graphiquement dans la figure 2. Espèces de basses altitudes ne dépassant pas 2.100 mètres environ (15) : Garrulax cinereifrons (150-1525), G. leucolophus (300-2.135), G. monileger (300-1.500), G. pectoralis (300-1.700), G. lugubris (800- 1.900), G. strepitans (900-1675), G. milleti (900-1.525), G. chinensis (niveau de la mer ou 200-1.375), G. nuchalis (300-800), G. vassali (100- 2135), G. galbanus (800 ou 500-1.830), G. delesserti (60-1.830), G. mitra- Source : MNHN. Paris DISTRIBUTION DES GENRES GARRULAX ET BABAX 151 tus (300-1575), G. ruficollis (300-2.000), G. canorus (niveau de la mer 1.830, et des plaines basses à 1.200 à Formose). Espèces s’étalant des basses aux hautes altitudes (4) : G. perspicil- latus (niveau de la mer-3.000), G. cineraceus (niveau de la mer ou 200-4.270). G. rufogularis (610-3.535), G. sannio (300-4.270). Espèces d’altitudes moyennes ne dépassant pas 3.000 mètres (14) : G. palliatus (1.220-2.230), G. rufifrons (915-2.500), G. striatus (800-2.740), G. lunulatus (1.500-2.500), G. caerulatus (1.065-2.750), G. merulinus (900- 2.440), G. cachinnans (1.220-2.650), G. jerdoni (1.070-2. ), G. virgatus (915-2. ), G. austeni (1.830-3.000), G. squamatus (1.070-3.000), G. yer- sini (1:800-2.500), G. formosus (900-3.000), G. milnei (1.375-2.800). Espèces d’altitudes moyennes dépassant 3.000 mètres (13) : Babar lanceolatus (1.400-4.850), Garrulaæ albogularis (1.220-4.575, et de 900 à 1.500 à Formose), G. maesi (800-3.650), G. variegatus (1.830-4.120), G. davidi (2550-4.000, mais nichant à environ 1.000 en Mongolie et Mandchourie), G. maximus (2165-4575), G. ocellatus (2.165-4.575), G. poecilorhynchus (1.500-4.575, et 1.200-2.100 à Formose), G. lineatus (:435-3.660, mais environ seulement 1.000-2.500 au Turkestan russe), G. subunicolor (1.830-4.880), G. elliotii (2.450-4.880), G. affinis (2.440- 4.880, et 2.000-3.000 à Formose), G. erythrocephalus (1.34 660). Espèces restreintes aux très hautes altitudes (5) : Babax waddelli (3.050-4.000), B. kKoslowi (3.900-4300), Garrulax sukatschewi (3.000- 4.000), G. bieti (3.050-4.575), G. henrici (2.930-4.575). TRAVAUX CITES Bancs, O., et Perens, J, L., 1928. — Birds collected by Dr. Joseph F. Rock in western Kansu and eastern Tibet. Bull. Mus. Comp. Zool. vol, 68, pp. 313-381. Benuioz, J., 1930. — Revision systématique du Genre Garrulax Lesson. Rev. d'Hist. Nat. part. 2, vol. 11, L'Oiseau, pp. 1-86. CmexG, T. H., 1958. — A distributional list of Chinese birds, Il, Passe- riformes. Peking. DeiGnan, H. G., 1945. — The birds of northern Thailand. Bull. U. Natl, Mus., no. 186. — 1964. — Timaliinae, in Check-list of birds of the world, vol. 10 pp. 240-427. Cambridge, Mass. 1933. — [Sans titre]. Bull. British Orn. Club., vol. 53, 8. DELAGOUR, J pp. 8: — 1946. — Les Timaliinés. L'Oiseau, vol. 16, pp. 7-37. Deracour, J., and JaBouizzx, P., 1931. — Les oiseaux de l’Indochine française, vol. 3, pp. 227-254. Paris. GreENwaY, J. C., Jr, 1933. — Birds from northwest Yunnan. Bull. Mus. Comp. Zool., vol. 74, pp. 109-168. Hacmisuxa, M., et Unacawa, T., 1951. — Contribution to the ornithology of Formosa, part. 2. Quarterly Jour. Taïwan Mus., vol. 4, pp. 1- 180. Hartent, E,, 1910. — The birds of Haïnan. Nov. Zool., vol. 17, pp. 189- Source : MNHN. Paris 152 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Lunzow, F., 1927-1928. — Birds of the Gyantse neighbourhood, southern Tibet. Ibis, 1927, pp. 644-659 ; 1928, 51-73, 211-232. — 1937. — In Lupcow, FE, et N. B. Kinwean. The birds of Bhutan and adjacent territories of Sikkim and Tibet. Ibis, pp. 1-46, 249-293, 467-504. _— 1944, — In Lunrow, F, et N. B. Kinnean. The birds of South- eastern Tibet. Ibis, pp. 43-86, 176-208, 348-389. — 1950. — The birds of Lhasa. Ibis, pp. 34-45. = 1951. — The birds of Kongbo and Pome, South-east Tibet. Ibis, pp. 547-570. Menenrzmacex, R, 1927. — Systematie results of birds collected at high altitudes in Ladak and Sikkim. bis, pp. 363-422, 571-633. __ 1998. —— Some biological problems connected with the Himalaya. Ibid, pp. 480-533. fuev, J. H., 1926. — A collection of birds from the provinces of Yun- nan and Szechwan, China, made for the National Geographic Society by Dr. Joseph F. Rock. Proc. U. S. Natl. Mus,, vol. 70, art, 5, pp. 1-70. _— 1931. — A second collection of birds from he provinces of Yun- nan and Szechwan, China, made for the National Geographic Society by Dr. Joseph F. Rock. Proc. U. S. Natl. Mus., Vol. 80, art, 7, pp. 1-91. Rwuey, S. D. 1961. — A synopsis of the birds of India and Pakistan. Bombay. Romwsow, H. C., 1928. — The birds of the Malay Peninsula, vol. 2, Lon- don. Suvrmes, B. E. 1953. — The birds of Burma. London. — 1960. — The birds of Borneo. London. Vaurue, C, 1959. — The birds of the Palearctic fauna, Passeriformes. London. Source : MNHN. Paris SPECIMENS OF LEAR’S MACAW IN THE ZOOLOGICAL MUSEUM OF AMSTERDAM by K. H. Voous Zoologisch Museum, Universiteit van Amsterdam The following remark by Sracer (1961) referring to the Machris Brazilian Expedition of 1956, reminded me of the presence of two unpublished specimens of Lear’s Macaw in the Zoological Museum of the University of Amsterdam : < A careful watch was kept for individuals of the rare Ano- dorhynchus leari, whose habitat is yet to be discovered, but no trace of the species was found in central Goiäs » (p. 15). In the old collection of mounted birds, which formerly belonged to the cabinet of the Amsterdam Zoological Gardens, I found five specimens of blue macaws, two belonging to the species hyacinthinus, one to glaucus, and two to leari. The. data follow below : Anodorhynchus hyacinthinus (Latham) Coll. Nr. 939. ©. « Brazil ». Wing 417, bill 87.5 mm. 940. SG. Zool. Garden Amsterdam, 7 April, 1884. Wing 438, bill 91.7 mm. Anodorhynchus glaucus (Vieillot) Coll. Nr. 941. G'. « Rio de la plata ». Wing 365, bill 68.2 mm. Anodorhynchus leari Bonaparte Coll. Nr. 942. « South America ». Wing 405, bill 67.1 mm. 943. Zool. Garden Amsterdam, 1882. Wing 391, bill 71.0 mm. Up to now no more than two specimens of À. leari were known to exist in collections : one being the type specimen in the Paris Museum (BONAPARTE, Naumannia, 6, 1856, Consp. Psitt., Beil. 1 ; Souancé, Iconographie des Perroquets, Paris, L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris 154 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE 1857, pl. 1) ; the other being a Zoological Garden (London) bird in the British Museum, Natural History (SALVADORI, Cat. Birds Brit. Mus., 20, 1891 : 149). The origin nor the exact dates, of these birds are known. Anodorhynchus leari is in coloration and size a wonderful intermediate between the well-known, large, Blue Macaw from central Brazil, and the equally well-known, smaller, Glaucous Macaw from southeastern Brazil, Paraguay, Uru- guay, and extreme northern Argentina. In the Amsterdam specimens of leari, which perfectly agree with Blanchard’s picture in SOUANCÉ (op. ci.), the upper parts are deep blue, but less intense than in hyacinthinus ; head, neck, and under parts have a decided greenish wash, the greenish tinge of the head contrasting remarkably with the blue of the mantle. Hyacinthinus is deep cobalt blue all over, and glaucus is greenish blue with a distinct greyish linge on the head and the neck, and a still darker grey wash over the feathers of the throat and upper breast. Apart from the specimens mentioned above captive Lear’s Macaws have been recorded to have been seen by BONAPARTE in the Antwerpen Zoo in the 1850’s (SouANCÉ, op. cit.) and to have been imported about 1925 from Parä to New York (fide Lee CRANDALL) and in the 1930’s to Germany (fide Oscar NEu- MANN ; PETERS, 1937 : 180, foot-note 1). In practically all of the scientific and popular works on parrots (including Cony, 1918; PErTErs, 1937 ; LEGENDRE, 1962) Lear’s Macaw is listed as a third species of the group of blue macaws. In view of the fact that the areas of A. hyacinthinus and A. glaucus are almost complementary, though probably more or less separated by the Brazilian Pla- teau, one would be anxious to know whether there are places where these species actually meet or overlap, and whether perhaps A. leari is no more than the result of a regular or sporadie, former or present, reproductive contact of hyacin- thinus and glaucus, and hence, in some way or another, a hybrid. The present scanty knowledge about this large and cons- picuous bird makes it unlikely, though not impossible, that it represents an independent species, of unknown geographic origin, intermediate between hyacinthinus and glaucus. The present author would feel highly rewarded, if this Source : MNHN. Paris LEAR’S MACAW IN MUSEUM OF AMSTERDAM 155 small note could stimulate someone to design accurate dis- tribution maps of the two well-known species of Anodorhyn- chus (which may prove a hard task !) and to show that these species have arisen through an almost classical way of geographic isolation in parts of Brazil, roughly north and south of the Brazilian Plateau. RESUME Partant des spécimens en peaux existant en collections, et des informations recueillies dans la littérature, l’auteur nous donne son point de vue sur la position systématique de l’Ara de Lear, qui est probablement une forme hybride de l’Ara bleu et l’Ara glauque, tout en souhaitant qu'une carte de dis- tribution précise des deux espèces bien connues d’Anodorhyn- chus puisse être bientôt établie. LITERATURE CITED Corx, G. B., 1918. — Catalogue of birds of the Americas. 2 (1). Publ. Field Mus. Nat. Hist. Zool. Ser., 13 (197). LeceNDre, M, 1962. — Perroquets et perruches. Paris (Boubée). Perens, J. L., 1937. — Check-List of birds of the world. 3. (Cambridge, Harvard Univ. Pr). Sracer, K., 1961 — The Machris Brazilian expedition, Ornithology Non-Pass. ines. Contr. Sc. Los Angeles Co. Mus., 41. Source : MNHN. Paris ADDITIONS TO THE LIST OF BIRDS OF THE REPUBLIC OF COLOMBIA by Alexander WETMORE Research Associate, Smithsonian Institution, Washington D.C. Président du X° Congrès International d'Ornothologie (Uppsala 1950) The observations that follow have come to attention during studies directed toward an account of the avifauna of the Isthmus of Panamä. In connection with these it has been necessary to review many species of birds that are found also in northern Colombia at the southern end of the great land bridge that connects the American continents. For this purpose collections have been assembled from little known areas in the Department of Chocé. It is appropriate to include these notes among those that honor M. Jacques BERLIOZ in recognition of his many contri- butions concerned with the birds of tropical America. A) DESCRIPTIONS OF NEW GEOGRAPHIC RACES Picumnus olivaceus malleolus subsp. nov. CHaracrers. — Similar to Picumnus olivaceus panamen- sis Ridgway of eastern Panamä (1), but darker above, more olive green ; breast and foreneck definitely browner ; streaks on under surface darker and heavier ; male with erown spots more yellow, less reddish orange ; slightly larger. DescriprIoN. — Holotype, U.S. National Museum no. 401870, male, La Raya, Bolivar, Colombia, collected Jan. 11, 1948, by M. A. CarrikeR, Jr. (orig. no. 12019). Crown and hindneck black ; pileum with small, elongated spots of () Picumnus olivaceus panamensis Ridgway, Proc. Biol. Soc. Washington, vol. 24, Feb. 24, 1911, p. 34. (Lion Hill Station, Panama Railroad, Canal Zone, Panamä). L'Oiseau et R.F.0., V. 35, 1965, n° spécial. Source : MNHN. Paris ADDITIONS TO THE BIRDS OF COLOMBIA 157 yellow ocher ; hindneck with scattered dots of white ; back, scapulars, and rump, light brownish olive ; wing coverts the same, margined very narrowly with chamois ; primaries and secondaries dusky with the outer web of the outer primaries edged narrowlÿ with buffy olive ; innermost primaries and secondaries bordered more widely with dull old gold ; upper tail coverts colonial buff ; rectrices black, the central pair with the inner web broadly yellowish white, the others tipped and margined with yellowish white ; side of head scantily feathered with dull white, the tiny feathers edged with black ; throat dull white ; foreneck and upper breast isabella color, somewhat paler centrally ; rest of under surface, including the under tail coverts, somewhat dull buffy white, streaked, rather narrowly, with dull olive ; under wing coverts light buff, mixed externally with brownish olive ; inner webs of primaries margined narrowly with buffy white. Bill dark neu- tral gray basally, black distally; tarsus, ioes, and claws fuscous black (in dried specimen). MEASUREMENTS. — Males (16 specimens), wing 50.6 - 55.7 (52.6), tail 23.4 - 26.6 (25.5 average of 15), culmen from base 11.1-12.6 (11.8, average of 15), tarsus 11.2- 13.3 (12.3) mm. Females (12 specimens), wing 51.8 - 53.8 (52.2), tail 24.0 - 26.7 (25.4), culmen from base 11.0 - 13.0 (11.8, average of 11), tarsus 11.4-13.5 (12.4) mm. RANGE. — Tropical lowlands of northern Colombia, in Cérdoba (Socarré and Quebrada Salvjin on the Rio Sinu), northern Antioquia (El Pescado, near Valdivia), and Bolivar (Regeneraciôn, La Raya, Volador, Santa Rosa, Colosé) in the drainage of the lower Rio Cauca and the lower Rio Magda- lena ; intergrades with Picumnus o. tachirensis east of the Rio Magdalena at Ocaña, Norte de Santander, and with P. o. olivaceus on the upper Rio Nechi, near Hacienda Belén above Zaragosa, northeastern Antioquia. Remarks. — The few specimens of this piculet previously available from northwestern Colombia, with an equally small number from Panamä east of the Canal Zone have been identified to the present as the typical race Picumnus oliva- ceus olivaceus. With the excellent series now at hand it is easily evident that Picumnus olivaceus panamensis Ridgway, named in 1911 from the Canal Zone, is a valid race, with a range extending through eastern Panamä, where it is recor- Source : MNHN. Paris 158 L’OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE ded on both Pacific and Caribbean slopes, to northern Chocé, Colombia (specimens from near Unguia). The population spread through northweslern Colombia, east of the Atrato drainage, represents another form that differs from panamen- sis as indicated in the description above. The typical race, P. o. olivaceus, is darker, more olive above, less definitely brown on the breast, with crown spots of the male decidedlÿ deep orange. At one of Carnixer’s localities, the Hacienda Belén, on a tributary of the Rio Nechi about 15 kilometers northwest of Segovia, he collected two males ; one, taken in the valley below the farm, with the head spots colored as in malleolus, but otherwise like olivaceus, and the other, from the hills above, that is typical olivaceus. The indication is that malleolus ranges up the Rio Nechi from its union with the Rio Cauca to near this point. These tiny woodpeckers in feeding peck so constantly and industriously, with as much noise as any of their larger rela- tives, that it seems appropriate to name this one malleolus, a small hammer. Heterospingus xanthopygius berliozi subsp. nov. CHARACTERS. Similar to Heterospingus æanthopygius æanthopygius (Selater) (1) but definitely darker : Male deci- dedly deeper black, with the yellow of lower back, rump, and lesser wing coverts darker ; female darker above, decidedly darker below, with the yellow of lower back and rump dar- ker. DescriprroN. — Holotype U. S. National Museum no. 443762, male, from Nuqui, Choc6, Colombia, collected March 5, 1951, by M. A. CarRiKER, Jr. (original num- ber 19777). Side of head and upper surface, except as noted, black ; superciliary stripe narrowly white immediately above eye, then, more broadly scarlet-red back to nape ; lesser wing coverts, lower back and rump between lemon chrome and lemon yellow ; foreneck and upper breast black, somewhat duller than the upper surface ; rest of lower surface fuscous- black : inner under wing coverts, axillars, and side of breast (1) Tachyphonus æanthopygius P. L. Sclater, Proc. Zool. Soc. London, pt 22, 1854 (April, 1855), p. 158, pl. 69 (Bogoté, Colombia). Source : MNHN. Paris ADDITIONS TO THE BIRDS OF COLOMBIA 159 (beneath the anterior end of the wing) white. Bill, tarsus, and feet black (from dried skin). MEASUREMENTS. — Males (6 specimens), wing 90.5 - 94.0 (92.1) ; tail 60.2-64.5 (62.9) ; culmen from base 18.4-19.6 (18.8) ; tarsus 21.7 - 22.5 (22.2) mm. Females (3 specimens), wing 86.2 - 92.2 (88.0) ; tail 63.8 - 65.4 (64.4); culmen from base 19.2-20.5 (19.9); tarsus 20.5 -21.8 (20.9) mm. RANGE. — Pacific slope of western Colombia from northern Chocé (Bahia Solano, Nuqui, Jurubidä) south to Caldas (Santa Cecilia) and Nariño (Guayacana). Remarks. — The definitely darker color of the series from which this form is described may be explained as a reflection of the greater rainfall of the coastal area of the northern Choc6. The series examined stands out clearly in comparison with 14 males and 7 females that cover the range of Hetero- spingus æanthopygius xanthopygius from eastern Darién in Panamä across northern Colombia to northern Santander. Those from the Rio Juradé, in northern Choc6, are interme- diate between the form here described and the nominale race, which is found on the Rio Jaqué, Darién, a short distance to the north. Skins in the Academy of Natural Sciences, avail- able through the kindness of R. M. de SCHAUENSEE, show that the subspecies berliozi ranges south over the Pacific slope to Nariño in southwestern Colombia. It is probable that this is also the bird of northwestern Ecuador. The name of this race is given to honor Jacques BERLIOZ in recognition of his contribution to knowledge of the avi- fauna of tropical America. It is offered as a token of our friendship through many years of association in researches in the field of ornithology. B) ADDITIONAL RECORDS FOR THE LIST OF BIRDS OF COLOMBIA The following notes cover several forms not reported pre- viously from the Republic of Colombia. The records are based on specimens that I have identified recently in collections in the U.S. National Museum. The names are presented in sys- tematic order. Source : MNHN. Paris 160 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Catoptrophorus semipalmatus semipalmatus (Gmelin). — R. M. SCHAUENENSEE in his recent book on The Birds of Colombia (1964, p. 83) in an account of the Willet lists the western subspecies, Catoptrophorus semipalmatus inornatus, with the remark that the typical form « probably occurs ». This supposition is verified by a female specimen in the U.S National Museum taken by M. A. CaRRIKER, Jr., at Coveñas, Bolivar, on Jan. 10, 1949. Measurements of wing 192.0 mm. exposed culmen 58.2 mm. and tarsus 55.8 mm., place this specimen with the smaller eastern subspecies. Actitis macularia rava Burleigh. — Thomas BURLEIGH in a recent review of the Spotted Sandpiper, Actitis macularia. has described a western race under the name À. m. rava, on the basis of lighter, grayer coloration above at all seasons, and a further minor distinction in a lesser amount of spotting on the lower surface in the plumage of the breeding season. Both the eastern and the western subspecies are represen- ted in the small series of Spotted Sandpipers in the Colom- bian collections made by CarrikEr for the U.S. National Museum. À male in winter dress, typical of the eastern race, A. m. macularia, was taken at Nuqui, Chocé, on Jan. 22, 1951. Specimens of À. m. rava, all in winter plumage, come from Unguia, Chocé, Feb. 28, 1950, Necocli, Antioquia, Jan. 26, 1950, and Coveñas, Bolivar, Jan. 13, 1949. Columba nigrirostris Sclater. — The Short-billed Pigeon ranges through the length of the Isthmus of Panamä so that it is not unexpected to find it in northwestern Colombia. Car- RIKER collected a male in heavy forest west of Acandi, Choc, Jan. 10, 1950. There is also a specimen in the American Museum of Natural History taken by H. E. ANrHony and D. S. Bazz March 29, 1915, on the eastern slope of Cerro Tacarcuna near the headwaters of the Rio Cuti, Choc6. In February and early March, 1964, I found these pigeons com- mon on Cerro Mali and the middle slopes of Cerro Tacarcuna in Darién adjacent to the Colombian boundary. It was evi- dent that they ranged throughout the forest that extended without a break across into adjacent Choc. Otus clarkit Kelso and Kelso. — During work in the mountains of eastern Darién, on March 3, 1964, Charles Source : MNHN. Paris ADDITIONS TO THE BIRDS OF COLOMBIA 161 O. Hanpzey, Jr. and I secured a male Bareshanked Screech Ovwl in a mist net set on a ridge of Cerro Tacarcuna, exactly on the line of the international boundary of the two republies of Panamä and Colombia. The slopes on both sides of the line are forested, with no distinction in habitat. The specimen is interpreted as a valid record for both countries, and on that basis is the first report of the species for Colombia. Several were recorded on the Darién side of the boundary. Glaucidium minutissimum minutissimum (Wied). — A male Least Pygmy Owl collected by M. A. CARRIRER, Jr. April 12, 1949, in virgin forest near the Rio Sin, at Socarré, Cérdoba, is the first record for Colombia. The place, called locally La Dispensa, is about two and one half kilometers below the mouth of the Rio Verde. The specimen agrees in color with those seen of the typical race from Brazil. Speotyto cunicularia brachyptera Richmond. — In 1941 in the Guajira, in northeastern Colombia M. A. CARRIKER, Jr. and I collected three Burrowing Owls of this race near Mai- cao on April 14, and one at Puerto Lépez, on April 22. These agree in pale coloration with specimens from northern Vene- zuela where this subspecies is widely distributed. Goldmania violiceps Nelson. — Goldman’s Hummingbird is common in the Serrania del Darién on the Panamanian side of the boundary on Cerro Mali, and over the slopes of Cerro Tacarcuna. On March 2, 1964 we caught one in a mist net set at 1460 meters elevation at the crest of the ridge on the boundary with Colombia. The occurrence of this species here has been overlooked, as there are two females and an imma- ture male in the American Museum of Natural History taken by H. E. ANrHonY on his expedition in 1915. The three birds, labelled « East slope of Mt. Tacarcuna, alt. 4600 feet, eastern Panamä », were collected April 10, 14, and 17, when the party was camped on the head of the Rio Cuti, Department of Chocé, Colombia. Picumnus olivaceus panamensis Ridgway. — Two speci- mens of the Olivaceous Piculet taken by CARRIKER March 1 and 18, 1950, at Unguia, Choc6 that agree fully with a series from eastern Panamä are the first report of this subspecies Source : MNHN. Paris 162 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE from Colombia. PErers in his Check-list (vol. 6, 1948, p. 90) placed panamensis as a synonym of olivaceus, In this he was misled by the scanty material that he had available, as birds from the Canal Zone through eastern Panamä are clearly dif- ferent in more yellowish brown shade above, and buffy gen- eral coloration below, with the breast distinctly paler. RESUME L'étude du matériel figurant dans les collections de l'U.S. National Museum a conduit l’auteur à décrire 2 formes nou- velles et à ajouter 8 espèces et sous-espèces à la liste des Oiseaux de Colombie. Source : MNHN. Paris A REVIEW OF THE SOUTHERN SUBSPECIES OF THE FAWN-COLOURED LARK MIRAFRA AFRICANOIDES SMITH by J. M. WiNTERBOTTOM Perey FitzPatrick Institute of African Ornithology, Cape Province When reporting on the Percy FitzPatrick Institute of Afri- can Ornithology - Windhoek State Museum expeditions to the Etosha Pan, I (WinrerBorroM, 1964) commented on the unsatisfactory state of the systematics of the Fawn-coloured Lark Mirafra africanoides Smith. The present paper is the outcome of that dissatisfaction. The Fawn-coloured Lark occurs in two discrete groups of populations, one in south-western Africa from the northern Cape Province Lo eastern Angola and western Zambia and the other in north-eastern Africa from northern Tanganyika to central Ethiopia and northern Somaliland. 1 make no attempt to deal with this second group and my remarks are confined to the southern populations. Within the range of these populations, 15 subspecies have been described, no fewer than seven of them from north- central South West Africa between 16° and 23°S and 16° and 19°E. The latest summaries available are those of McLacHLAN & LivERsipGE (1957) and Wuire (1961); and MACDONALD (1957) has commented on the South West Afri- can races. MeLacHLan & LIVERSIDGE recognise nine subspecies (one further subspecies is extra-limital to their work), sup- pressing one (M. a. omaruru (Roberts) recognised by Mac- DONALD ; White récognises eight subspecies, with another three as too poorly defined to be worth maintaining. The subspecies recognised by both these authorities (and by Mac- DONALD if they oceur within the area with which he was con- cernéd) are : L'Oiseau et R.F.O., V. 35, 1965, n° spécial. Source : MNHN. Paris 164 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE a. africanoides Smith, a. harei Roberts, a. sarwensis (Roberts), . a. austinrobertsi White, a. vincenti (Roberts), a. makarikari (Roberts), a. rubidior White. MeLacHLan & LiVERSIDGE and MAcponaLD both recognise also M. a. gobabisensis (Roberts), which WuHiTE considers only separable at the 30 % level from M. a. africanoïdes ; and Wire also recognises M. a. trapnelli White, which is extra-limital for the others. He suppresses M. a. mossam- biquensis Pinto, recognised by McLacHLaN & LIVERSIDGE, as a synonym of M. a. vincenti. All three authorities are agreed that M. a. isseli HOESCH & NIETHAMMER is not a valid form and Wuite also suppresses M. a. quaesita Clancey, which was described subsequent to the publications of the others (but see below). None recognise M. a. ovambensis (Roberts) or M. a. tsumebensis (Roberts) but whereas MacpoNaLD fuses both with M. a. sarwensis, White puts the first with sarwensis as a poorly defined variation and the second with M. a. africanoïdes as an intergrade bet- ween that and sarwensis. Finally, there is a nomenclatorial problem involving the use of the names africanoides, austinrobertsi and quaesita. Wuire (1956) quotes personal information from MACDONALD that Surrw's types of M. a. africanoïdes do not agree with birds from Litakun, near Kuruman, which place ROBERTS had proposed as the type locality, but do agree with speci- mens from Colesberg, which Wuire (1961) considers are aus- tinrobertsi. CLANCEY (1958) points out that this finding means that austinrobertsi must be sunk in the synonymy of africa- noides and has named the Kuruman birds quaesila ; but this depends on the identity of birds from Colesberg and from Nylstrom (the type locality of austinrobertsi), on which see below. Enough has now been said, I hope, to reveal the differences of view. I have had at my disposal 478 skins and I must thank the following for the loan of material : The Director of the Durban Museum & Art Gallery (Mr. P. A. CLANCEY) ; the Director of the East London Museum (Miss M. Cour- TENAY-LATIMER) and Mr. C. D. QuickELBERGE ; the Director Source : MNHN. Paris eh - MNHN Paris Fig. 1. — Map of Southern Africa, showing ranges of subspecies of Mirafra africanoides. Hatched : Areas of intergradation and/or 166 L'OISEAU ET LA REVUE FRANÇAISE D’ORNITHOLOGIE and Mr. M. P. Sruart IrwIN of the National Museum, Bula- wayo ; the Director and Mr. P. J. Buys of the State Museum, Windhoek ; the Director and Mr. O. P. M. Prozesky of the Transvaal Museum, Pretoria ; Mr. J. Huzc of the University Museum, Oxford ; and Mr. H. von MALTZAHN. Variation in the Fawn-coloured Lark is considerable in colour, from deep red-brown in rubidior to grey and buff in makarikari ; and in the degree of dark slriping above ; and small in the extent of spotting below and in size. Colour changes and changes in the extent of striping are rarely abrupt ; and there is considerable overlap in the ranges of some of the subspecies, though whether this is due to local migration, to nomadism, to ecological factors or to some other cause we have not enough data to determine ; but the presence of many non-breeding specimens of M. a. sarwensis from the Tsotsoroga Pan, 140 miles north of Maun, is pretty clear proof that considerable movements oceur. As CLANCEY has pointed out (see above), the finding that Smrrw’s types agree with Colesberg, rather than Kuruman birds leaves the Kuruman and other northern Cape birds without a name ; and he accordingly named them quaesila, type locality Riet- fontein, Asbestos Mountains. Many of the birds from this area are richer, redder and less heavily streaked than afri- canoïides but less so than gobabisensis (Roberts), which occu- pies the area to the north-west. However, if their range is plotted on a map, it is found to fall almost entirely within the ranges of the other two races and quaesita is best discarded as an intergrade between africanoides and gobabisensis ; though a small area round the type locality appears to be populated solely by such birds. The total number of subspe- cies recognised below is 10, two more than Ware (1961) reco- gnises for the same area ; but four of these are not very well defined and could be suppressed without much distortion of the picture. The races are as follows : 1. MIRAFRA AFRICANOIDES AFRICANOIDES Smith Mirafra africanoides A. Smith, Rept. Exp. Expl. C. Afr, 1836 : 47 - Eastern province of the colony and Latakoo (res- tricted type locality, Colesberg, C. P.). A dark race, heavily streaked above on a dark reddish brown ground. Source : MNHN. Paris SOUTHERN SUBSPECIES OF MIRAFRA AFRICANOIDES 167 Range. — The north-eastern Cape, from Colesberg and Prieska to Vryburg, and the adjacent parts of the Transvaal and Orange Free State. Intergrades with gobabisensis over a wide area of the northern Cape. Measurements. — 34 Gÿ : Wing 88-97 mm, av. 92.5 ; tail 53-63, av. 58.7 ; culmen, 15-17, av. 16.2 ; tarsus 18-24, av. 21.4. 4 99 : Wing 85-89 mm ; av. 87.2 ; tail 51-57, av. 53.7 ; culmen 15-16, av. 15.7 ; tarsus 19-22, av. 20.7. Material examined. — Durban Museum, 9 (Kuruman, Riverton, Prieska) ; East London Museum, 20 (Griquatown, Kimberley, Barkly West, Kuruman, 25 m. N. of Coles- berg) ; Transvaal Museum, 7 (Vryburg, Bloemhof, Fourteen Streams) ; South African Museum, 1 (Colesberg) ; National Museum, 1 (30 m. E. of Prieska). Total, 38. 2. MIRAFRA AFRICANOIDES AUSTINROBERTSI White Mirafra africanoides austinrobertsi White, Ibis, 1947 : 420 (nom. nov. for Anacorys africanoïdes transvaalensis Roberts, Ann. Tol. Mus., 18, 1936 : 262 - Nylstroom ; nec Mirafra afri- cana transvaalensis Hartert, 1900). Inseparable in appearance from M. a. africanoides but smaller. Males are 85 % separable according to whether the wings are 90 mm and over or 89 mm and under. Mr. A. N. Rowan who kindly analysed the figures, tells me that the difference is statistically highly significant. Range. — The northern Transvaal and western Rhodesia, north to the western Gwaai Reserve, where it meets {he next form. Measurements. — 8 G : Wing 85-93 mm, av. 89.9 ; tail 45-58, av. 53.1 ; culmen 15-17, av. 16.0 ; tarsus 18-20, av. 20.4. 2 99 : Wing 84-87 mm ; tail 46-49 ; culmen 15-16 ; tar- sus 19-22. Material examined. — National Museum, 6 (Chatsworth, Bulawayo, Coplestone, Gwaai Reserve, Nyamandhlovu) ; Transvaal Museum, 5 (Border of Bushveld, Olifants River, Warmbath). Total, 11. Source : MNHN. Paris 168 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE 3. MIRAFRA AFRICANOIDES VINCENTI (Roberts) Anacorys africanoides vincenti Roberts, Ostrich, 1938 : 117 - Umvuma. Synonym : Mirafra africanoides mossambiquensis Pinto, Bol. Soc. Est. Mocamb., 22, 1952 : 5 - Maquese, Alto Chan- gawe. Another rather poorly defined form, but the ground colour above is paler and yellower than austinrobertsi and the strea- king not quite so heavy above ; more speckling on the throat. Range. — Rhodesia, from Wankie to Umvuma and Maran- dellas ; and southern Portuguese East Africa. Intergrades with austinrobertsi at Copleston and Nyamandhlovu. Measurements. — 19 Gœ : Wing 85-94 mm, av. 90.3 ; tail 48-57, av. 53.4; culmen 14-18, av. 16.1; tarsus 18-23, av. 21.0. 5 99 : Wing 78-86 mm, av. 84.0 ; tail 46-53, av. 50.0 ; culmen 15-16, av. 15.4; tarsus 19-23, av. 21.6. Material examined. — National Museum, 12 (Coplestone, Wankie, Gwaai Reserve, Selukwe, Marandellas, Funhalanso) ; Transvaal Museum, 4 (Umvuma, Alto Changane) ; Durban Museum, 9 (Panda, Manhica). Total, 25. 4. MIiRAFRA AFRICANOIDES TRAPNELLI White Mirafra africanoides trapnelti White, Bull. B.O.C., 64, 1943 : 21 - Between the Kasisi and Litapi Rivers, Balovale, Zambia. Very similar to the last two forms but rather paler and averaging larger. Range. — Western Barotseland, from Sesheke to Balovale, but not the extreme west, where maharikari replaces it. Measurements. — 7 a : Wing 89-95 mm, av. 91.9 ; tail 46-58, av. 54.5: culmen 15-17, av. 16.1; larsus 19-21, av. 20.6. 1 © : Wing 84 mm ; tail 50 ; eulmen 15 ; tarsus 22. Material examined. — National Museum, 8 (Balovale, Loma, Chiolola). Total, 8. Source : MNHN. Paris SOUTHERN SUBSPECIES OF MIRAFRA AFRICANOIDES 169 5. MIRAFRA AFRICANOIDES MAKARIKARI (Roberts) Anacorys africanoides makarikari Roberts, Ann. Tvl. Mus., 15, 1932 : 28 - Nkate, Bechuanaland Protectorate. A grey form, with buff edges to the feathers above ; deci- dedly paler than any of the preceding subspecies. Range. — The north-eastern Kalahari, north to the wes- tern Sesheke and Senanga Districts of Zambia, where it meets trapnelli. Overlaps in the south-west of its range, and again north-east of Maun, with sarwensis ; the second of these areas of overlap being probably due to off-season movements of sarwensis. Measurements. — 30 Go : Wing 84-97 mm, av. 89.3 ; tail 45-61, av. 53.8 ; culmen 14-17, av. 15.9 ; tarsus 19-23, av. 21.3. 13 90 : Wing 82-88 mm, av. 85.4 ; tail 48-57, av. 51.6 ; culmen 14-17, av. 15.5 ; tarsus 18-24, av. 20.6. Material examined. — National Museum, 34 (Nata, 61 m. W. of Nata, 32 and 40 m. N. of Nata, Nangweshi, Bushman Pits, 70 m. E. of Maun, 80 m. N. of Maun, Tsotsoroga Pan, Ghanzi, Chiolola, 25 m. N. of Shangombo, Luete, Senanga, Nkate Road, Tsane) ; Transvaal Museum, 11 (Dekar, 30 m. S. W. of Sehitwa, Cungona, Tsumkwe). Total, 45. 6. MIRAFRA AFRICANOIDES SARWENSIS (Roberts) Anacorys africanoïdes sarwensis Roberts, Ann. Toi. Mus., 15, 1932 : 27 - Kaotwe Pan, Bechuanaland Protectorate. Synonym : Anacorys africanoides ovambensis Roberts, Ostrich, 1937 : 97-50 m. N. W. of Namutoni. Redder and darker than makarikari. Range. — The central and north-western Kalahari, west across northern South West Africa to about 15°E. Overlaps with omaruru on the Okavango and the south-east corner of the Etosha Pan; and with gobabisensis in western Bechua- naland between 20° and 21°S. and 21° and 23°E. Measurements. — 60 G' : Wing 84-97 mm, av. 91.1; tail 49-62, av. 55.5; culmen 14-18, av. 15.8 ; tarsus 18-24, av. 21.1. Source : MNHN. Paris 170 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE 25 9 9 : Wing 84-96 mm, av. 88.1 ; tail 48-59, av. 54.3 ; culmen 13-18, av. 15.3 ; tarsus 18-23, av. 20.6. Material examined. — National Museum, 111 (Hukantsi, Inkowane Pan, Kikomu di Kaï, 18, 57 and 70 m. east of Kakia, 41 m. W. of Kanye, Kukong, Kofi Pan, Letlaking- Mashaweng road, Lothlekane, Lehututu, Lephepe, Lake Dow, Mache, Murumush, Molepole-Lephepe road, 80 m. N. of Maun, Odiakwe, Okwa, Sekhuma Pan, Serowe-Inkowane Pan road, 13 m. N. W. of Serowe, Tsabong and 63 m. N., Tsepe, Tsot- soroga Pan, Tshane) ; Transvaal Museum, 35 (Xobi, Tsane, Hukantsi, Tierputs, 20 and 95 m. S. of Ghanzi, Kobi Pan, Dekar, Drotzky, 30 m. S. W. of Seketwa, Aha Mis, Bohelo- labaha, Damara Pan, Gemsbok Pan, Tsumkwe, Kaotwe, Kuke) ; South African Museum, 5 (Namutoni, Onoolongo, Onguma) ; State Museum, 1 (Onguma) ; Okavango Expedi- tion, 4 (Runtu, Kapaku, Makamba, Omuramba Omutaka). Total, 156. 7. MIRAFRA AFRICANOIDES HAREI Roberts Mirafra africanoïdes harei Roberts, Ann. Tol. Mus., 5, 1917 : 258 - Windhoek. Synonym : Mirafra africanoides isseli Hoesch & Nietham- mer, J. f. Orn., Suppl., 1940 : 215 - Okahandja. More heavily streaked and richer red than gobabisensis. Range. — Central South West Africa from Otavi Lo Kal- krand, where it meets gobabisensis. Overlaps with omaruru in the Outjo area. Occasional birds from as far east as Barkly West and Francistown are indistinguishable. Measurements. — 19 G'œ : Wing 87-96 mm, av. 914; tail 49-60, av. 55.1; eulmen 14-17, av. 15.6 ; tarsus 19-23, av. 21.0 14 99 : Wing 85-92 mm, av. 87.8 ; tail 51-57, av. 53.3 ; culmen 13-17, av. 14.9; tarsus 18-23, av. 20.7. Material examined. — Transvaal Museum, 15 (Windhoek, Okahandja, 22 m. E. of Dordabis, Tsumis, Quickborn, 40 m. W. of War Grave, Barkly West, Akangunde, Outjo, Otjiwa- rongo, Maltahohe Road) ; National Museum, 7 (20 m. S. of Letlaking, 15 m. S. of Tsabong, 22 m. E. of Molepole, 92 m. S. of Francistown, Tshane, 45 m. E. of Kakia) ; East London Source : MNHN. Paris SOUTHERN SUBSPECIES OF MIRAFRA AFRICANOIDES 171 Museum, 3 (Windhoek, Kalkrand) ; State Museum, 5 (Win- dhoek, Otjivarongo) ; Durban Museum, 4 (Windhoek, Oka- handja), von Malizahn collection, 4 (Otavi). Total, 38. 8. MIRAFRA AFRICANOIDES GOBABISENSIS (Roberts) Anacorys africanoides gobabisensis Roberts, Ann. Tol. Mus., 18, 1936 : 263 - Gobabis. Decidedly less heavily streaked above than harei; paler, redder and less heavily streaked above than africanoides. Range. — The southern Kalahari, from Gobabis to Oli- fantshoek and Upington. Intergrades freely with africanoides over a wide area of the northern Cape ; and overlaps sarwen- sis in southern Bechuanaland Protectorate. Measurements. — 39 G'œ : Wing 86-98 mm, a. 93.2 ; tail 48-62, av. 56.0 ; culmen 14-17, av. 15.7 ; tarsus 17-24, av. 21.3. 15 9 9 : Wing 86-89 mm, av. 86.4; tail 50-57, av. 54.2 ; culmen 13-16, av. 14.9 ; tarsus 19-23, av. 21.2, Material examined. — National Museum, 27 (Tsabong, 63 m. N. of Tsabong, Kukong, Kakia, Tsume Kikomu di Kai, 20 m. W. of Kanye, Lokwabe, Lehututu, Murumusa Pan, Gobabis, Nkane) ; Transvaal Museum, 10 (Gobabis, 25 m. E. and 20 m. W. of Upington, Kalahari Desert, Bohepabata, Aub- Nossop confluence) ; East London Museum, 20 (Tsumis, Wit- sand, Eindpaal, Kuruman-Askham, Teekoms, Olifantshoek, Klipput) ; South African Museum, 4 (Morokwen, Upington, Griqualand-West) ; Durban Museum, 5 (Lidfontein, Prieska, Kuruman, Mata-Mata). Total, 67. The details of the intermediates between M. a. africanoides and M. a. gobabisensis, which CLANGEY calls quaesila, are as follows : Measurements. — 28 GS : Wing 87-98 mm, av. 925 ; tail 51-62, av. 56.4; culmen 15-19, av. 15.9; tarsus 18-95, av. 21.2. 15 9 Q : Wing 84-95 mm, av. 88.5 ; tail 48-60, av. 54.5 ; culmen 14-17, av. 15.7 ; tarsus 18-23, av. 20.6. Material examined. — Durban Museum, 14 (Rietfontein (type), Riverton, Kuruman) ; East London Museum, 18 (Mat- Source : MNHN. Paris 172 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE zap-Volop, Klipput, Witsand, Kalahari Game Reserve, Wit- draai) ; South African Museum, 5 (Witdraai, Modder River) ; National Museum, 5 (Riverton, Kimberley, Witdraai) ; Trans- vaal Museum, 1 (Niekerkshoop). Total, 43. 9. MIRAFRA AFRICANOIDES OMARURU (Roberts) Anacorys africanoides omaruru Roberts, Ann. Tol. Mus., 18, 1936 : 263 - Omaruru. Synonym : Anacorys africanoïdes tsumebensis Roberts, Ostrich, 1937 : 98 - Tsumeb. Less heavily streaked and yellower above than hareï ; paler and more richly coloured than sarwensis. Range. Northern South West Africa, overlapping sarwensis on the Okavango and eastern Etosha Pan and with harei in the Outjo area. Measurements. — 18 Sœ : Wing 85-98 mm, av. 91.9; tail 48-61, av. 54.3; eulmen 14-18, av. 15.3; tarsus 19-23, av. 20.9. 6 99 : Wing 84-94 mm, av. 89.1; tail 52-53, av. 52.7; culmen 15-17, av. 15.8 ; tarsus 20-22, av. 21.0. Material examined. — South African Museum, 9 (Oma- ruru (type), Onguma, 60 and 75 m. W. of Okakuejo, Otji- vasandu, Kowares, Kameeldoorn waterhole) ; Transvaal Museum, 10 (Outjo, Orupembe, Ohopoho, Kowares, 30 m. N. of Omaruru) ; Okavango Expedition, 4 (Rooïdag, Makamba, Omaramba Omutaka) ; National Museum, 1 (35 m. W. of Kang); State Museum, 1 (Kowares) ; von Malizahn coll., 1 (Erongo). Total, 26. 10. MIRAFRA AFRICANOIDES RUBIDIOR White Mirafra africanoides rubidior White, Bull. B.O.C., 75, 1955 : 29 - Ozondache, west of the Waterberg. Heavily streaked on a rich, reddish-brown ground, much deeper and richer than in any other form. Range. — Only known from the type locality, and its range is entirely surround by that of harei. Source : MNHN. Paris SOUTHERN SUBSPECIES OF MIRAFRA AFRICANOIDES 173 Measurements. — 2 Gœ : Wing 95 mm; tail 60, 63; culmen 17 ; tarsus 22. 1 @ : Wing 87 mm; tail 58 ; culmen 13 ; tarsus 20. Material examined. — Oxford University Museum, 3 (Ozondache (type)). Total, 3. If we plot male and female average wing-lengths, the subspecies fall into two groups, a larger (africanoides, rubi- dior, gobabisensis, haréi, sarwensis and omaruru), in which male wing-lengths average from 91.4 to 95.0 mm and female D *RUBIDOR 294 > î < eGOBABISENSIS QUE) ô ï FRICANODS a decrrat 5 92 LTRAPNELLI +OMARURU 2 *HAREI … eSARWENSIS 2 à PVINCENTI Le 2AUSTINROBERTSI MAKARIKARI 89 #4 tr] 30 65 86 87 88 AV, WING-LENGTH OF FEMALES (AM) Fig. 2. — Average wing-lengths of male and female of subspecies of Mirafra africanoides. from 86.4 to 89.1 ; and a smaller (trapnelli, vincenti, austin- robertsi and makarikari) in which the figure for male and female are 89.3-91.9 and 84.0-88.1 respectively. Only in males of trapnelli is there any overlap between the groups, of which the larger is distributed in the south and west and the smal- ler in the north and east. Source : MNHN. Paris 174 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE RESUME Révision, basée sur 478 peaux, des différentes sous-espèces jusqu'ici décrites de Mirafra africanoides, desquelles 10 seu- lement sont retenues, encore que 4 d’entre elles (marquées d’un astérisque) peuvent être discutées M. a. africanoides Smith, M. a. austinrobertsi White *, M. a. vincenti (Roberts) *, M. a. trapnelli White *, M. a. makarikari (Roberts), M. a. sarwensis (Roberts), M. a. harei Roberts, M. a. gobabisensis (Roberts), M. a. rubidior White, M. a. omaruru (Roberts) *. REFERENCES Crancex, P. A. (1958). — Miscellaneous Taxonomie Notes on African Birds, X, 1. Durb. Mus. Novit., 5 : 99-101. Macpowaz», J. D. (1957). — A Contribution to the Ornithology of Wes- tern South Africa. London. McLacnLan & Livensiner (1957). — Roberts Birds of South Africa. Johannesburg. Wavre, C. M. N. (1956). — Notes on African Larks, Part. II. Bull. Brit. Orn. Club, 76 : 53-60. Wavre, C. M. N. (1961). — A Revised Check List of African Broadbills, Pittas, Larks, Swallows, Wagtails and Pipits. Lusaka. Wivrensorrom, J. M. (1964). — Report on the Birds of Game Reserve No. 2, Cimbebasia, 9. Source : MNHN. Paris Supplément à L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Volume 36. — 1966, n° 1 TABLE DES MATIÈRES Volume 35. — Nouvelle Série. — 1965 1 — Numéros 1 et 2 (Voir plus loin la Table du N° Spécial qui complète les publications de l'année) TABLE ALPHABETIQUE DES AUTEURS DONT LES ARTICLES ET LES NOTES (*) SONT PUBLIÉS DANS CES FASCICULES Arrerr (0). — Note sur trois oiseaux migrateurs non encore signalés à Madagascar en Arrerr (0). — Découverte de la nidification de Puffinus pacificus (Gmelin) près de la côte ouest de Madagascar .. BenLioz (J.). — Note critique sur les Trochilidés lia et Angasma 1 Biscnorr (M.) et Srnpen (J. C.). — Observation d’un Tichodrome échelette (Tichodroma muraria) dans les Vosges moyennes .... 154 BLaise (M.). — Contribution à l'étude de la reproduction du Cou- cou gris Cuculus canorus dans le nord-est de la France at87 BLonDeL (J.). — Le Héron cendré Ardea cinerea L. nicheur en Camargue .. 59 Cuisin (M). — L'imitation des cris du Pie noir par certains T didés é 155 Decacour (JL). — La Sare en Nouvelle-Calédonie 66 ErarD (Ch) et Meniner (D. d'Oléron 22 Fonmox (A.). — Récentes données sur la nidification de Asio flam- meus (Pontop.) en Bourgogne et en Forez ......,.,,........ 63 Fourier (0.) et Jarry (G.). — Observations et capture de Cal- carius lapponicus en Vendée ...... 54187 Founwien (O.) et Jannx (G.). — Capture d' ie des-moissons de Sibérie (Anser fabalis rossicus Buturlin) en Vendée ........., 158 Genatn (M). — Observations ornithologiques en Algérie oceiden- tale 46, 117 Janny (G.). — Quelques observations et captures intéressantes en Seine-et-Marne ............ 69 Janny (G.). — Voir Fournier (0.) 157, 158 Source : MNHN. Paris 168 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE Lazrrre (A.) +. — Recherches sur la durée de la période de repro- duction chez quelques oiseaux d’Eure-et-Loir de 1920 à 1962. + Mauzx (P). — Aperçu faunistique sur l'ile Juan-de-Nova 152 * Mansaz (L). — Nidiñeation du Hibou grand-due Bubo bu Roussillon .... 60 Menucen (D). — Voir Eranp (Ch.) 22 Prévost (1). — Note sur le baguage des oiseaux de i'arehipel de Pointe Géologie de 1950 à 1963 ; 17 * Srnouer (J. G.). — Voir Biscnorr 154 VisuuanD QD. — Aspect de l'avifaune ouessantine en octobre novembre . 140 Vincent (F.). — Le Coucon émeraude (Chrysococcyx cupreus inter- medius Hartlaub 1857) parasite d'un nid de Souimanga (Nec- tariniidae) au Congo (Brazzaville) 81 wassesiom (V. — Répartition de la Pie-griche à poitrine rose (Lanius minor) en Catalogne du nord-est . 151 Yearax (L). — Les spécimens de migrateurs paléarctiques en pro- venanee d'Afrique tropicale dans les collections du Muséum. 27 TA re TT Te Ce en 70 TABLE ALPHABÉTIQUE DES SUJE TRAITÉS DANS CES FASCICULES Anas aucklandica chlorotis en Nouvelle-C Anser fabalis rossiens en Vendée Ardea cinerea nicheur en Camargue Asio flammeus nicheur en Bourgogne et Angasma, note critique sur le genre a Baguage des oiseaux à lArchipel de Pointe Géologie : Bourgogne, nidification du Hibou brachyote 63 Bubo bubo nicheur en Roussillon . 60 Calcarius lapponieus en Vendée 157 Camargue, reproduction du Héron 59 Catalogne, répartition de la Pie-grièche à poitrine rose 151 Chrysococcyx cupreus intermedins parasite un Souimanga s1 Collections de migrateurs paléaretiques d'Afrique tropicale au Muséum : 27 Congo, parasite 81 Cuculus canorus, reproduetion 87 Forez, nidification du Hibou brachyote 63 Imitation des cris du Pie noir par certains Turdidés 155 Juan da Nova, aperçu faunistique ........... 152 Lanius minor, répartition en Catalogne du nord-est 151 Madagascar, migrateurs nouveaux ... ï 66 Madagascar, nidifieation de Puffinus pacifieus .. 135 Migrateurs paléarctiques d'Afrique tropicale au Muséum 27 Nectariniidae parasité par le Coucou émeraude ....... 81 Nouvelle-Calédonie, présence de la Sarcelle brune 66 Oléron, observations hivernales 22 Ouessant, avifaune en octobre-novembre 140 Période de reproduction d'oiseaux d'Eure-et-Loir 9 Pointe-Géologie, baguage des oiseaux ... Ë 17 Puffinus pacificus, nidification à Madagascar . 135 Reproduction, recherches sur la durée en Eure-et-Loir . 9 Source : MNHN. Paris L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE 169 Reproduction de Cuculus canorus 87 Roussillon, nidification du Grand-Due . 60 Seine-et-Marne, observations et captures intéressantes 69 Tichodroma muraria dans les Vosges moyennes 154 Timolia, note critique sur le genre 1 Trochilidés, systématique 1 Turdidés imitant le Pie noir 155 Vosges, observation d'un Tichodrome 154 BIBLIOGRAPHIE OUVRAGES Aux (S). — The Birds of Sikkim 71 Axpensen (T), Josnsex (4. H), Nonnsvane (A), Panensex (E. oi) Preuss (N. D.). — Danmarks Fugle . ; 71 Arrison-Wizces (G. L). — Wildfowl in Great 72 Bere (F. Lo et Wasmn GW); North American Falconry and Hunting Hawks 160 CLacæy (P. A). — The Birds of Natal and Zululand 161 Friscx (0, von). — Bei sellenen Vôgel in Moore und Steppe ...... 161 IMMELMANN (Dr K). — Die australischen Plattschweifsittiche ..... 73 Ivaxov (A. I.) et Srecman (B. K). — Clef de détermination des oiseaux Œ'U.R.S.S. 73 Jacourmann (8). — Les derniers rapaces . 4 Jonnson (N. K). — Biosystematies of sibling species of flycat- chers in the Empidonaz hammondii - oberholseri - wright COPIE RO + ë 162 LanpsRonouGH THOMSON (À 7 Mexzez (H). — Der Steinschmätzer ........ T6 Niërmammer (G), Kramer (H.) et Wozrens (H. E.). Deutschland 77 No (H). — Die Brutvôgel in ihren Lebensgebicten 162 ParzoLn (R). — Die Feldlerche .......... 76 Presrwrem (A. A). — I name this parrot 77 Russeuz (S. M). — A distributional study of the birds of British Honduras 78 Scnrrr (R früher 78 SCHONWETTER 163 : [4 Surrmens (Re HN) A Check-List of the Blrds of the Bechua naland Protectorate and the Caprivi Strip Svexsson (L.). — Bestämni Tams (T. J. C). — De vogels van Terschelling, overzicht van alle op het eiland waargenomen sorten 78 Turrs (R. W.). — The Birds of Nova-Scotia 79 Vaurie (C). — The Birds of the Palearctie Fauna ST RE RC lee 165 Warcux (D: Ph. de). — Les oiseaux de cage et de volière . 79 Divers Le problème des oiseaux sur les aérodromes ................... 166 Verslag van het vergelijkend onderzock naar de wuze van vœdsel- zocken van enige soorten steltlopers ........................ 80 Source : MNHN. Paris 170 L’OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE TABLE DES ILLUSTRATIONS Cartes de répartition des captures de migrateurs paléarctiques en Afrique tropicale figurant dans les collections.du Muséum .... Côte sud-ouest de Madagascar . Poussin et Biotope de nidification de Bubo bubo en Rou @L D ee 62 Secteur Domjevin-Fréménil (M.-et-M.) 100 Vallée de la Meurthe et ses affluents .. 97 Source : MNHN. Paris TABLE DES MATIÈRES Volume 35. — Nouvelle Série. — 1965 II — Ne spéeial Le Professeur Berlioz, Volume jubilaire TABLE ALPHABETIQUE DONT LES ARTICLES SONT PUBLIÉS DANS CE FASCICULE Amanon (D.). — Notes on the Galapagos Hawk . 9 BANNERMAN (D). — Some reflections on two recent visits to the Azores archipelago ; 22 Bounuiène (F.) et Ércmécorar (R. D.). — Introduction : 1 Broëxnuysex (G. J.). — An analysis of bird casualties on the roads in the South Western Cape Province, South Africa FRS Deracour (J.). — Les Pigeons de la Nouvelle-Calédonie | 52 Demenriev (G. P.) et. SrÉPAnyaN (L. S.). — Quelques particularités de la reproduction des oiseaux dans les hautes zones du Thian- Chan . 58 Donsr (JL). — Le Professeur Berlioz 3 Gwer (A). — Sur le Faisan de Hume et le genre Calop) 65 Hônsrantus (S.). — De l'effet désastreux de l'enrobage des semences sur l’avifaune suédoise nl Laëx (D.) et Moreau (R. E). — Clutch-size in tropical passerine birds of forest and savanna . 76 Man (E). — Classification, identification and sequence of genera and species 90 Nigruammer (G.). — Zur Taxonomie von Cinelus cinclus. Ein tier- geographisches Problem : 96 Rupcey (S. D). — Le Martinet pâle de Socotra 101 SaLomowsex (F.). — Notes on the Sunbird-Asitys (Neodrepanis) .. 103 Smuex (C. G.). — Molecular systematies : New techniques applied to old problems 112 Srreseman (E.) et (V. chus pugnaz 125 THomsox (A. L. Shearwater . 130 VaumE (C.). — Distribntion régionale et altitudinale des genres Garrulax et Babaz et notes sur leur systématique .......... 141 Voous (N. H). — Specimens of Lear's Macaw in the Zoological Museum of Amsterdam 153 Wermone (A). — Additions to the list of birds of the Republic of Colombia 156 Winrernorrom (J. M.). — A review of the southern subspecies of the Fawn-coloured Lark Mirafra africanoides Smith .......... 163 Source : MNHN. Paris 176 L'OISEAU ET LA REVUE FRANÇAISE D'ORNITHOLOGIE TABLE ALPHABETIQUE DES SUJETS TRAITÉS DANS CE FASCICULE Açores (observations ornithologiques) .... 22 Anodorhynchus leari (spécimens du Museum d’Am: 153 Apus pallidus berliozi de Socotra ...... 101 Babax (distribution, notes systématiques) 141 Biologie de la reproduction de la Buse des 9 Buteo galapagoensis (notes sur) 9 Calophasis (notes sur le genre) 65 Cinelus cinelus (taxonomie) 96 Classification (généralités) $ . 90 Colombie (additions à l'inventaire ornithologiq 156 Distribution altitudinale et régionale des genres Garrulaæ e ou die da om 141 Enrobage des semences et son effet désastreux en Suède . 7 Faisan de Hume ... : es 65 Garrulax (distribution, notes systématiques) . 141 Migrations transéquatoriales du Puffin des Anglais 130 Mirafra africanoides (revue des races méridionales) 163 Mortalité des oiseaux due à la cireulation routière 35 Mue du Chevalier combattant ................ 125 Neodrepanis (notes systématiques) 103 Nouvelle-Calédonie (Pigeons) 52 Philomachus pugnaz (mue) 125 Pigeons de la Nouvelle-Calédonie . Ve 52 Pontes des passereaux des forêts et Savanes tropicales 76 Puffinus puffinus (migrations transéquatoriales) 130 Reproduction des oiseaux du Thian-Chan . 58 Socotra (Martinet pâle) te 101 Systématique moléculaire 112 Taxonomie de Cinelus cinelus . 96 Taxonomie de Mirafra africanoides cure 163 Thian-Chan (particularités de la reproduction des 58 TABLE DES ILLUSTRATIONS Le Professeur Jacques Beniioz à sa table de travail (PI. D . 1 Le Professeur Bencroz photographiant des Proteas (PI. I) 8 Passer melanurus, Telephorus zeylonus, Streptopelia senegalensis, Streptopelia capensis (PI. IT) Ne RE Bubo africanus, Lanius collaris, Ploceus capensis, Serinus cani- collis (PL IV) ar, 39 Distribution des genres Garrulax el Baba . 145 Distribution des races de Mirafra africanoides . 165 ERRATUM N° spécial 1965, article G. Nisrmammen, p. 100, 4° ligne du RÉSUMÉ : au lieu de : e type cinclus à ventre pâle », lire : « type cinelus à ventre foncé ». Source : MNHN. Paris