THE QUATERNARY AVIFAUNA OF CRETE, GREECE by Peter n.M. WEESIE* TABLE OF CONTENTS Page Abstract, Resume ............ ................... .................. ................... .......... .............. ......... .... 3 I. 1. Introduction ........................................................................................ 4 2. Acknowledgements .............................................................................. 5 11. Localities, Material, Age, Methods, Terminology and Abbreviations ............................ 6 I. Localities .......................................................................................... 6 2. Material .............. .............................................................. ....... ......... 7 3. Age .......................... ,', ...... ,"','" .. , ............ " .. ,"""""'"'''''''''''''''''''''' 7 4. Methods ................... ,", .. " ...... " ............ , .. , ... , ...... , .... , ....... ,., ... ,""""'" 8 4,1, Identification ..... , ................................. , ....................... ",., .............. , .. , 8 4,2, Statistics ... , ... ,. """ , .... , .... , ............ ,' .. ,", ... ,', .. ,', ............ ,.,', ... ,., ... "...... 8 5, Terminology " ............ ,", ...... , ......... , ......... ,""""', ................... ,""'"'''' 9 6, Abbreviations """" ...... " ... ", ... , ... """ ..... " ....... ,.,' .,,""",., "" .. "" ... " .. ," ID 6, 1. Osteological abbreviations """" .. "." ........ """"" .. """" .. " .... ",, .. ,,""",,. 10 6,2. Abbreviations of museum names ........ " .... " ...... " .... """ .... " ... ",,............ ID 6.3 Abbreviations of localities """"" ........ " .... "" .. " .. """"" ... ,,"""""""",,. 10 • Institute for Earth Sciences, State University Utrecht, Budapestlaan 4, 3508 TA Utrecht, Netherlands, Key-words: Avifauna, Quaternary, Crete, Systematics Mots-c1es : Avifaune, Quaternaire, Crete, Systematique Palaeover/ebra/a, MantpelJier, 18 (I): 1-94,9 figs" 30 tab., 9 pis. (Re~u le 2 Juin 1987, acceptc le 3 Septembre 1987. public le I"' Septembre 1988) 2 III. Systematics ....... ............ ...... ....... ...... ............ ....... ......................... ...... ...... .......... 11 Order Procellariiformes .... ...... .......... .......... ....... ...... .......... ....... ...... .......... ........ 11 Order Pelecaniformes ...................................................................................... 12 Order Anseriformes ........................................................................................ 12 Order Accipitriformes ..................................................................................... 13 Order Galliformes .......................................................................................... 24 Order Gruiformes ......... ...... ................. ........... ........ .............. ........ ...... ............ 25 Order Charadriiformes ...... ....... ......... .......... ..... ....... .......... ...... ................ ......... 26 Order Columbiformes ..................................................................................... 27 Order Strigiformes.......................................................................................... 30 Order Caprimulgiformes.................................................................................. 33 Order Apodiformes ........ ..................... ......... .............. ....... ...... ....................... 34 Order Piciformes............................................................................................ 35 Order Passeriformes........................................................................................ 35 IV. The localities and their species contents .................................................................. 51 1. General remarks ... ......... ....... ........ ......... .................... ......................... 5 I 2. Number of identified species and their fossil remains ................................. 51 3. Species association from Liko cave ........................................................ 52 4. Species association from the other localities ............................................ 54 V. Comparison of the Pleistocene and Recent avifaunas and the paleoenvironment on Pleistocene Crete ......................................... :...... ...... .......... ...... ..................... 54 I. General remarks ...... .................... .............. ........................ .......... ....... 54 2. Comparison of the composition of the Pleistocene and Recent avifaunas .......................................................................................... 54 3. Comparison of U,e climatic zones inhabited by the species of U,e Pleistocene and Recent avifaunas ........................................................... 56 4. The habitats of the species and the paleoenvironment ................................ 57 VI. Characteristics of the Pleistocene cretan avifauna. a basis for discussion and speculation ...................... ...................................................... ...................... ..... 57 1.1 The climate and surface area fluctuations of Pleistocene Crete ..................... 58 1.2 The paleobiogeography of some birdspecies in relation to the climate and vegetation on Pleistocene Crete ............ .......... .......... .......... ............. 58 2. The absence of most Galliformes .......................................................... 59 3. The abundance of birds of prey .. ...... ...... ............................................... 60 4. The endemic birds of prey................................................................... 60 VII. Some remarks on Island avifaunas and the speciation of birds of prey on Islands ......... .................... ............................... ............................. ............... 62 VllI. Disappearance and extinction of various Pleistocene cretan bird species ....................... 63 References ........ ......................... ........ ............................ ..................... .................... 64 Tables .................................................................................................................... 69 Captions of Plates ..... ...................... ............................. ......... ........ ........................... 90 To the memory of my sister Karin A reconstruction of some Pleistocene Cretan birds: Gyps melitensis, Aqui/a chrysaetos simurgh, Athene cretensis and Garrulus glandarius. 3 ABSTRACf Pleistocene bird fossils have been studied from nine localities on Crete. Part of this material was described earlier by the auulOr (Weesie, 1982) and will not be treated here in extenso, the results will be incorporated. More than one third of the over 10,000 fossil bird bones available could be identified; they were found to represent at least 65 bird species. The following species of the Pleistocene Cretan avifauna are new to the fauna of Crete: Branta nificollis, Haliaeetus albicilla, Gyps melitensis, Aquila chrysaetos simurgh n. ssp., Ketupa zeylomensis, Aegoliusfimereus, Dendrocopos leucotos, Zoothera dauma, Turdus iliacus and Pyrrhula pyrrhula. The Pleistocene Cretan avifauna differs less from comparable mainland avifaunas than (fossil) avifaunas fram oceanic islands do. Still, the Pleistocene Cretan avifauna has two qualities that are characteristic of island avifaunas : the almost complete absence of a group of birds (the Galliformes) and the presence of two endemic (sub)species : the giant eagle Aquila chrysaetos simurgh n. ssp. and the long-legged owl Athene cretensis (Weesie, 1982). The new subspecies is described in the present study. These endemic birds of prey were found in association with their supposedly principal prey species (now extinct as well) : endemic mice for the owl and endemic deer for the eagle. Endemic mammals have been found in association with endemic birds of prey on many islands, not only in the Mediterranean. There is evidence that the size of endemic birds of prey becomes optimally adapted to their feeding on certain endemic mammals, especially rodents. Another characteristic of the Pleistocene Cretan avifauna is the great number of species of birds of prey. This appears to be a common characteristic of fossil avifaunas from caves on Mediterranean islands as well as from caves on the European mainland. However, we think that ecological conditions on Pleistocene Crete (especially the abundant presence of mice) helped to account for the high representation of birds of prey. Furthermore, the fossil avifauna enables us to draw some conclusions about the climate and vegetation on Pleistocene Crete: it is concluded that the climate was cooler than today and that Crete was largely covered with forests. Finally, the reasons for the extinction or disappearance from Crete of some bird species of the Pleistocene Cretan avifauna are discussed. Des fossiles Pleistocenes d'oiseaux de neuf localites en Crete ont ete etudies. Une partie de ce materiel a ete dccrite deja par l'auteur (Weesie, 1982) et ne sera pas traitee ici extensivement, les resultats serant incorpores dans le present travail. Plus d'un tiers de la quantite de plus de 10,000 ossements disponibles pouvail etre identific et au moins 65 eSpCces d'oiseaux sont representces dans cet ensemble. Les especes suivantes de l'avifaune Pteistocene de Crete sont nouvelles pour la faune de Crete: Branta ruficollis, Haliaeetus albicilla, Gyps melitensis, Aquila chrysaetos simurgh n. ssp., Ketupa zeylonensis,