Biology and Conservation of the Andalusian Buttonquail (Turnix sylvaticus sylvaticus Desf. 1789) Carlos Gutiérrez Expósito PhD 2019 A mi familia A Raquel “Almost throughout the higher sections of the animal kingdom you have males fighting for the females, the females caring for the young; here, in one insignificant little group of tiny birds, you have the ladies fighting duels to preserve the chastity of their husbands, and these latter sitting meekly in the nursery and tending the young. It is, to our ideas, a very odd arrangement, because we have become so thoroughly imbued with the spirit of the opposite one; but it answers apparently just as well, so far as the interests of the race are concerned, as that one with which we are so familiar, and on which we pin our faith; and in this and many similar cases it has often seemed to me that nature mutely warned mankind against dogmatism and against the foolish, though all too prevalent, belief that only what we know and are used to can be good, and that neither government nor society can get along equally well under any laws and forms but just those which we are become accustomed.” Allan O. Hume & Charles H.T. Marshall 1879 The Game Birds of India, Burmah, and Ceylon Recommended citation: Gutiérrez‐Expósito, C. (2019) Biology and Conservation of the Andalusian Buttonquail (Turnix sylvaticus sylvaticus, Desf. 1789). PhD Thesis. Universidad Pablo de Olavide. Sevilla ‐ Estación Biológica de Doñana (CSIC). This PhD project has been funded by the grant BES‐2015‐074938 of the Severo Ochoa support program to Centres of Excelence in R + D + I (SEV‐2012‐0262‐01) of Ministerio de Ciencia, Innovación y Universidades of the Spanish Government and has been conducted in the Conservation Biology Department at Estación Biológica de Doñana (CSIC). Additional economical support has been obtained from SEO/BirdLife, Autoroutes du Maroc and Espacio Natural Doñana. All fieldwork in Morocco has been done with the collaboration of the Institut Scientifique ‐ Université Mohammed V de Rabat, under the authorisation of the Haut Commissariat aux Eaux et Forêts et à la Lutte Contre la Desertification du Royaume du Maroc. © All pictures by the author. © Cover illustration by José Antonio Sencianes (diariodeunacuarelista.blogspot.com). Any form of reproduction, distribution, public communication or transformation of copyrighted material is forbidden without explicit permission by the author. To accomplish with the standards of the Severo Ochoa program for Centres of Excellence, this PhD project has an international advisory committee with researchers of recognized international prestige in the area: Dra. Beatriz Arroyo Research Scientist Game and Wildlife Management Research Group Instituto de Investigación en Recursos Cinegéticos Ciudad Real (Spain) Dr. Philip McGowan Professor of Conservation Science and Policy School of Natural and Environmental Sciences University of New Castle New Castle upon Tyne (United Kingdom) Dr. Pedro Jordano Research Professor Department of Integrative Ecology Estación Biológica de Doñana (CSIC) Sevilla (Spain) Biology and Conservation of the Andalusian Buttonquail (Turnix sylvaticus sylvaticus Desf. 1789) Carlos Gutiérrez Expósito PhD ‐ Sevilla, 2019 Table of contents Abstract ............................................................................................................................. 1 Resumen ............................................................................................................................ 7 General introduction ....................................................................................................... 13 Objectives ........................................................................................................................ 25 Methods and techniques ................................................................................................. 29 Section 1: The Turnicidae family ..................................................................................... 41 Chapter I Who’s first? An index to priorize monitoring and conservation efforts of the poorly known buttonquails (Turnicidae)......................................................... 43 Chapter II Asymmetric heterochromia in birds: the dark crescent of buttonquails ........ 67 Section 2: Biology and conservation of the Andalusian Buttonquail .............................. 75 Chapter III Vanishing wildlife in populated areas: the demise of the Andalusian Buttonquail ...................................................................................................... 77 Chapter IV The farmland refuge of the last Andalusian Buttonquail population .............. 93 Chapter V Breeding ecology of the Andalusian Buttonquail (Turnix sylvaticus sylvaticus) ....................................................................................................... 115 Chapter VI Gardens by the sea: the last Andalusian Buttonquail refuge ........................ 126 General discussion ......................................................................................................... 129 Findings / Conclusiones ................................................................................................. 135 Supplementary material ................................................................................................ 139 References ..................................................................................................................... 171 Agradecimientos ............................................................................................................ 199 ________________________________________________________________ Abstract Abstract 1 Abstract _________________________________________________________________ Abstract Rise of agriculture and livestock growing during the Mesolithic‐Neolithic transition gave birth to a new biotope, the farmland, which in a few thousand years reached a planetary scale at the expense of natural ecosystems. This widespread land use change implied a demise of many avian species as its natural habitats were replaced by crops and grazing lands. However, it became into a new habitat for some steppe, grassland and savannah birds who were able to shift from its natural habitats to human‐made environments. Some avian species of many bird families found there a land of opportunity, among them a few buttonquail (Turnicidae) species. However, the recent farmland intensification is leading to a general decline of these species. In this PhD we use the Andalusian Buttonquail (Turnix sylvaticus sylvaticus) as a case study of a taxon, at least in part, able to cope with traditional farming, which until recently had a wide distribution in the western Mediterranean countries, and currently is in the verge of extinction. Starting from a global and historical scale approach, contextualizing the taxon at biogeographic and taxonomic levels, we will arrive at the "in situ" study of the only population that still survives. Firstly, we will try to establish the historical evolution of the extinction process in Europe and North Africa, to model and evaluate the historical distribution area based on environmental variables and to establish the phylogeographic relationships of the historical populations of this subspecies and their relationship with the rest of the species of the family. Secondly, the study of the population present in Morocco is carried out in order to determine the size of the population at different times, to evaluate the population trend and to describe habitat selection and reproductive biology. Chapter I. Who’s first? A rarity index to prioritize monitoring and conservation efforts of the poorly known buttonquails (Turnicidae). Wildlife conservation status assessments are usually made at species level. However, the risk of extinction of subspecies or local populations often go unnoticed due to the lack 2 ________________________________________________________________ Abstract of a proper assessment. Buttonquails are very secretive birds whose conservation assessment has proven to be very difficult due to the lack of good field data. Here we used the Turnicidae family to create a rarity index, which can also be applied at subspecific level, based exclusively on data provided by eBird, a citizen science birding platform. At species level, our rarity index matches with the IUCN red list categories assigned to every buttonquail species, classifying at higher level the most endangered or unknown species: New Caledonian Buttonquail (Turnix novaecaledoniae), Luzon Buttonquail (Turnix worcesteri) and Buff‐breasted Buttonquail (Turnix olivii). By applying this index at subspecific level, we found that at least 12 buttonquail subspecies have a rarity index similar or higher than these three buttonquail species, all of them being island endemisms of Common Buttonquail (Turnix sylvaticus), Barred Buttonquail (Turnix suscitator) and Red‐backed Buttonquail (Turnix maculosus) from the Philippines, Indonesia and Papua New Guinea. We propose the use of this index to detect bird species, subspecies or populations which could be in high risk of extinction and are lacking the necessary field studies. Chapter II. Asymmetric heterochromia in birds: the dark crescent of buttonquails. I describe for the first time the unique coloured pattern of the iris of buttonquails (Turnicidae). This unique pattern is due to the presence of a dark‐brown crescent in the iris below the pupil, whose form and extent varies