Friedhelm Weick Owls (Strigiformes) · Annotated and Illustrated Checklist Eulen (Strigiformes) · Kommentierte und illustrierte Artenliste Friedhelm Weick Owls (Strigiformes) Annotated and Illustrated Checklist With 63 Illustrations and 23 Color Plates Friedhelm Weick Pommernstraße 34 76646 Bruchsal-Untergrombach Germany About the Author Friedhelm Weick was born in Karlsruhe, Germany. For 35 years he worked as a technician in the construction industry. Throughout this time he painted birds, and since 1965 he has also been a professional illustrator and wildlife artist. Since 1988 he has been a graphic designer at the Museum for Natural History in Karlsruhe, a job which entailed painting a complete range of animals and plants. He illustrated over 120 books, including most vol- umes of the Handbuch der Vögel Mitteleuropas (1971–1997) and all volumes of Die Vögel Baden- Württembergs (1987–2006). His work has appeared in all leading wildlife magazines. In addition, he has published books in his own right, including Birds of Prey of the World, in collaboration with L.H. Brown, Owls, a Guide to the Owls of the World, in collaboration with C. König and J.H. Becking and Anmut im Federkleid, in collaboration with O. Kröher. Library of Congress Control Number: 2006927285 ISBN-10 3-540-35234-1 Springer Berlin Heidelberg New York ISBN-13 978-3-540-35234-1 Springer Berlin Heidelberg New York This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illus- trations, recitations, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer. Violations are liable to prosecution under the German Copyright Law. Springer is a part of Springer Science+Business Media springer.com © Springer-Verlag Berlin Heidelberg 2006 Printed in Germany The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Editor: Dr. Dieter Czeschlik, Heidelberg, Germany Desk Editor: Dr. Jutta Lindenborn, Heidelberg, Germany Cover design: design&production, Heidelberg, Germany Typesetting: Stasch, Bayreuth, Germany Production: LE-TEX Jelonek, Schmidt & Vöckler GbR, Leipzig, Germany Printing and binding: Stürtz AG, Würzburg, Germany Printed on acid-free paper 31/3100 YL – 5 4 3 2 1 Dedicated to three splendid owl-experts and friends / Den drei hervorragenden Eulenkennern und Freunden gewidmet: Dr. h. c. Siegfried Eck †, Dresden, Germany Dr. Gerrit Paulus Hekstra, Harich, The Netherlands Professor Dr. Claus König, Ludwigsburg, Germany Foreword The owl is a magical creature: it exudes an ambivalent fascination that can trigger completely antithetical responses. On the one hand, there is something unsettling about the owl’s soundless flight, eerie call, and nocturnal activity that can result in its demonization as a messenger of darkness. On the other hand, however, its masklike visage and seemingly penetrating gaze prove en- thralling, and this may have contributed to the deification of the owl as the companion spirit of an omniscient mother earth. In addition, the highly spe- cialized captor of prey aroused the competitive envy of hunting parties, even provoking a bit of resentment when they lost out to owls in their hunt for small game. Moreover, the excrement and pellets (hairballs) they deposited in barns and church towers contributed to their reputation as undesirables or even pests. With the overcoming of superstition and suspicion, our relation- ship with these creatures of the night has now become one of unreserved ad- miration. They grant us insights into the rich diversity of adaptation to noc- turnal hunting as well as into sound inventories, brooding biology, and nest- ling development. In a complete transformation of what was once a symbol of fear and hatred, charismatic owls now enjoy an almost singular popularity in large parts of the population. Whether it be the barn owl, eagle owl, or boreal (Tengmalm’s) owl, each is well-suited as the flagship species for a biotope and species protection in keeping with our times. A gratifyingly large number of specialists, ama- teurs, and enthusiasts are involved in owl protection: creating potential nest- ing hollows, banding birds, collecting pellets, guarding endangered breeding areas, or mitigating the risk of electrocution by power poles. The organization of owl friends on the local, regional, and international level ensures that advi- sory support and coordination take place at a high level of expertise. Other friends of owls seek to regenerate owl populations in as natural a way as possible, whether for resettlement or for securing the continued existence of highly endangered species. This effort encompasses not only zoos, bird parks, and special breeding stations, but also those work-intensive rehabilitation cen- ters in which injured or orphaned owls can be given professional care. All of these organizations and facilities serve, directly or indirectly, to inform and enlighten the public and are thus instrumental in ensuring a positive image for the once so detested “wraiths of the night.” The spectrum of owl-related hobbies includes nature photography, painting and sculpture, with the lattermost encompassing the most diverse forms of owl representation. Owls it appears are highly reluctant to let go of anyone who has fallen into their clutches. viii Foreword This certainly applies to the author of this taxonomic list of owl species. Deeply impressed by the grace and patterns of behavior of wild animals, Friedhelm Weick moved from a very different profession to become – about 40 years ago – a painter of animals and an illustrator of textbooks. The begin- nings of his collections of illustrations and sources from the literature, as well as his study of museum skins (to obtain an overview of the world of owls) go back at least 25 years. These activities were considerably boosted by his joint efforts with Claus König to create the first compendium of the entire order of Strigiformes (1999). Such tenacious and long-term pursuit of this project un- doubtedly reflects a profound enthusiasm for owls. In a time of ever-new media that bombard our senses and constantly seek to outdo each other in emotional impact, that dazzle us with floods of images, 3-D projections, and virtual reconstructions of long-lost worlds, a systematic list of a specific group of birds may seem anachronistic and antiquated. And, in fact, there is no lack of voices that ridicule the outmoded museum work of classical taxonomists and see no practical value in discussions of the system- atic classification of organisms. Since academia too did not remain untinged by this subliminal disdain, the foundations of species information and sys- tematic classification were largely forced out of research and teaching in the competitive struggles among the various academic disciplines. If the univer- sity, however, shirks its responsibility, taxonomy threatens to become an en- dangered discipline. And this despite the fact that sound knowledge of the systematic subdifferentiation of organisms represents the essential precondi- tion for understanding evolutionary processes. In a multifaceted interplay with geography, ecology, ethology, and genetics, systematics makes it possible to trace the natural history of populations. This includes the origin and paths of dissemination of populations, their development through competition and predation, through changes in climate or shifts in available habitat, and their isolation and radiation into distinctive forms of adaptation. Ever since the environmental protection conference in Rio de Janeiro in 1992 (the “Earth Summit”), the catch-all term of “biodiversity” has enjoyed pervasive use. Even if biodiversity extends the variety of biological phenom- ena to habitats and landscapes, there is no doubt that diversity of organisms continues to represent its most important foundation. This lends new weight to the genealogies of organisms. The majority of developed nations signed the Biodiversity Treaty, which makes mandatory the protection of the worldwide diversity of lifeforms. Within the European Union, the concept of “Natura 2000” is based on this objective. Aside from the “Red Lists” of endangered species, criteria were also established for Germany for determining responsibility in the protection of species. Even if these are highly commendable developments, they nevertheless remain first and foremost a reflection of a troubling process of extinction, an ever-accelerating disappearance of species caused by a his- torically unprecedented expansion of human settlement. Whether projects of research involve questions of evolutionary biology, con- servation programs for the preservation of the species, or the breeding of eco- nomically relevant livestock, knowledge and correct taxonomy of “species” represent the unconditional precondition for all of them. Even if the naming and categorizing of animate objects is as old as the development of human language itself, the scientifically grounded definition and differentiation of particular groups of organisms as a “species” is nonetheless a highly complex Foreword ix discipline. And here, apart from objectifiable decision-making criteria, sub- jective aspects such as worldview, system of social values, or even national ambitions and political correctness may have a part to play. Moreover, the state of knowledge regarding global diversity and its diffusion patterns remains full of gaps. According to present listings we know of 1.7 million recent species on earth from the realms of protozoa, fungi, plants, and animals. Approximately 50000 of them are vertebrates, of which 9935 are birds; a good 200 of these, in turn, are owls. However, since many of the tropical forests have not yet been ex- plored and may possibly be destroyed by logging operations prior to a species survey, well-founded speculations reckon with the possibility of up to ten times as many species. Early museum taxonomy was only concerned with classificatory systems for designating and administering the diversity of organisms, and this was especially true as long as even biologists considered the immutability of cre- ation as incontrovertible dogma. Ever since Darwin, however, the goal has be- come more ambitious: to classify species as the basis for reconstructing their evolutionary history (phylogenesis). A comparative examination of morpho- logical, physiological, and ethological properties that takes the results of ecol- ogy and zoogeography into account expects, on the one hand, that all indi- viduals of a population demonstrating a high level of matching traits can be brought together as of the same species and, on the other hand, that a high level of matching traits between heterogeneous populations correlates with a high degree of kinship between them. Ernst Mayr added, as an essential crite- rion of differentiation of individual species, the reproductive barrier develop- ing between them in evolution, which minimizes the risk of interspecies hy- bridization. Nevertheless, all classic species concepts are confronted by an abun- dance of convergent forms, which is why they have to place great weight on the distinction between homologous and analogous traits. Additionally, a com- parison of traits is frequently insufficient for the systematic differentiation of subspecies and even less sufficient for assessing higher-level taxa in terms of their phylogenetic connections. Ultimately, molecular biology, at the end of the 20th century, has led the way out of this stagnation on the systematic level in an almost revolutionary man- ner, for it turned out that very fundamental questions could only be approached by incorporating modern genetics into the analysis. Above all the structural analysis of genetic material, of DNA, has revived the discussion of taxonomic classification and given it back the societal recognition it needs. Since DNA analysis looks for similarities in genetic material, it is more immune to irrita- tions arising from convergences in morphology. The goal here goes beyond the grouping of related species; the effort is to reconstruct as realistically as possible the process of speciation with its individual branchings of the phylo- genetic tree. It is even possible to locate such splitting off points in time with the help of the “molecular clock,” which makes the phylogenetic tree into a multidimensional construct. Of all the groups of organisms, the phylogenetic classification of birds is the most advanced. Currently, recent forms have been arranged into 23 orders, 146 families, and 2141 genera of 9935 species (as of December 2005, according to zoonomen.net). Spectacular fossil discoveries of recent years, especially from mesozoic deposits in China, even allow the documentation of the derivation of x Foreword birds from feathered, bipedal saurians (of the Sauriurae and Ornithurae fami- lies). These fossils, of at least 120 million years in age, can already be assigned to the plovers, loons, petrels, and anserines. Owls have existed for 50–60 million years. Within the class of birds, owls represent a conspicuously uniform morpho- logical group. With their usually large and rigidly forward oriented eyes within the “face” of a large, round, and “neckless” head, with their loose, fleecy plumage usually in camouflaging colors, and with their broad and compact bodies, it is not hard for anyone to recognize them as owls. Insofar, however, as the uniform ap- pearance of owls was selected by a nocturnally active lifeform as predator, this uniformity of morphological traits made systematic subdifferentiation more dif- ficult. Given this unity, there was never any controversy regarding the owls as a uniform group, even though as early as 1840 Nitsch had proposed splitting them up into a group of barn owls and a group of “typical owls.” There was, however, a great deal of uncertainty regarding their integration with their next of kin, as Sibley and Ahlquist (1990) showed in great detail in their historical survey. On the basis of their similar beak forms, Linnaeus (1758) grouped owls, as the genus Strix, together with falcons and shrikes (in the Accipitres order). Baird (1858) was the first to use Strigidae as a family name for nocturnal birds of prey, in the context of the bird world of North America. Huxley (1867) clas- sified owls with vultures and secretary birds. Fürbringer (1888) was the first to dissociate owls from diurnal raptors, incorporating the Striges lineage with the family Strigidae among the Coraciiformes. Peters (1931) was the first to recognize owls as an independent order Striges, ranking them among night- jars, parrots, and cuckoos. According to Sibley and Ahlquist, the methods of molecular biology made it possible to partially corroborate these kinship relationships. Nonetheless, in this interpretation, owls were separated from the Coraciiformes and at the same time moved closer to the swifts and hummingbirds. DNA analysis requires the dissociation, once and for all, of nocturnal from diurnal birds of prey. These authors add a new superorder, Strigimorphae, which includes the Strigiformes order, with its 319 species in 51 genera. A suborder, Strigi, within the latter, includes the two owl families, Tytonidae, with its 15 species in 2 genera, and Strigidae, with its 184 species in 27 genera. As of late, the frogmouths and the nightjars are also incorporated as suborders within the Strigiformes order. Within the Strigimorphae superorder, the Strigiformes order is joined by the turacos (Musophagiformes order). The systematic list of the owls of the world provided here encompasses 220 species of owls and with the subspecies classification results in 539 taxa. By providing synonyms, the aim is to clear up the often confusing duplication found in the designations. (Even today, valid species names are sometimes assigned twice: Glaucidium passerinum, for instance, designates not only the Eurasian pygmy owl, but also a blue-blossomed plant in Japan.) This checklist has been brought up-to-date and already takes the results of genetic analysis into account. This has led to the splitting up of previous species complexes, as is especially striking in the case of the genera Otus and Glaucidium. In this context, Weick names considerably more owl species than do all authors be- fore him: Peters (1940) cited 141 species, del Hoyo et al. (1999) cited 195, the World Birds Taxonomic List (zoonomen.net) cites 199, and König et al. (1999) cited 212. Recently, however, the Global Owl Project actually named 225 owl species (www.globalowlproject.com/species). The current upheaval in owl sys- Foreword xi tematics is documented by countless individual publications. By bringing to- gether sources that are in part widely dispersed, the succinctly annotated check- list provided here presents a kind of status report. It goes without saying that no list can ever be final, since the process of bringing the classification into line with the findings of evolutionary biology represent a continuous process of approximation and convergence. On the one hand, animal forms exist in a continuum of superspecies, species, and subspe- cies, of ecotype and morph, and in this way the construct of the species con- cept ultimately originates from a convention, which has no equivalent in real population dynamics. Thus, in spite of all scientific differentiation criteria, there remains a large amount of leeway for personal assessments. On the other hand, the temporal and spatial differentiation of the branchings in the phylo- genetic tree is much more multifaceted than can be reproduced through the limitations of rigid classificatory categories (Newton 2003). Furthermore, evo- lutionary biology has demonstrated that in the course of evolutionary history a cooperative fusioning of heterogeneous organisms must have repeatedly oc- curred, which means that individuals at a higher level of organization actually have emerged from “species conglomerates,” which raises a problem of philo- sophical theory for the definition of species. This checklist is neither an identification guide, nor a mere picture book, nor an owl monograph. By reflecting a high level of scientific expertise and the most up-to-date research, it provides outstanding support in the handling of scientific collections and in studies of global biodiversity. But it should also address the needs of practitioners of scientific species preservation, since global, national, or regional responsibility for ensuring the continuance of endangered forms fre- quently first becomes apparent on the basis of taxonomic and geographic dif- ferentiation. Not least of all, this list can support scientific conservation and breeding programs, whose contribution to species preservation is largely de- pendent on knowledge of the systematic subclassification of the animals bred. I too would like to support this intention to the best of my abilities. St. Oswald, Germany January 2006 Dr. Wolfgang Scherzinger Zoologist in Bavarian Forest National Park
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