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Long-term trends, total numbers and species richness of increasing waterbird populations at sites on the edge of their winter range: cold-weather refuge sites are more important than protected sites

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Abstract

Recent climate changes are most likely the major determining factor for the increasing importance of Central Europe for wintering waterbirds, given that most of the region is located on the edge of these species’ wintering ranges. A few recent large-scale studies have demonstrated changes in species distribution at the flyway level, but detailed studies at the site level are still scarce. Using mid-January wintering waterbird counts from 532 sites taken over a 48-year period (1966–2013), we have assessed the role of ‘cold-weather refuges’, i.e. sites where the selective pressures of winter harshness are reduced (e.g. sites with ample sources of running water, urban areas, warmer sites, sites with a relatively higher proportion of wetlands), and a site’s protection status on species richness, total numbers and trends at the site level. We found prevailing increasing trends in total numbers and species richness at the site level, which are in line with the area’s increasing importance as wintering grounds. However, some sites have likely been affected by density-dependent regulation as mean total numbers per site have not increased since the 1990s. Density dependence may also be a reflection of an increasing number of sites with running water in traditionally cold areas. Factors affecting trends in species richness at the site level are however less predictable. We demonstrate the great importance of cold-weather refuges, where running water has an effect on total numbers, species richness and trend affect total numbers, warmer areas and higher proportions of wetlands in the surroundings affect total numbers and species richness and urban areas affect total numbers. However, we found that legislative safeguards for sites, such as the establishment of Nature Reserves and Special Protection Areas, have no significant effects. Our findings therefore suggest that the effect of temperature and presence of cold-weather refuges (as defined here) are more relevant to this observed increasing trend of wintering waterbirds on the edge of wintering ranges than a reduction in human developmental pressures.

Zusammenfassung

Langfristige Entwicklungen, absolute Individuenzahlen und Artenreichtum wachsender Wasservögel-Populationen am Rand ihrer Winterquartiere: vor kaltem Wetter geschützte Quartiere sind wichtiger als solche in Schutzgebieten

Die Klimaveränderungen der letzten Zeit sind vermutlich der Grund für Zentraleuropas wachsende Bedeutung als Winterquartier von Wasservögeln; schließlich liegt Zentraleuropa zumeist am Rand der traditionellen Winterquartiere dieser Arten. Obwohl einige neuere Groß-Untersuchungen gezeigt haben, wie sich die Verteilung der Arten entlang der Zugrouten verändert, weiß man auf der Quartiers-Ebene über solche Veränderungen nur wenig. Unserer Untersuchung lagen jeweils Mitte Januar durchgeführte Wasservögel-Zählungen an 532 Standorten und über 48 Jahre (1966–2013) hinweg zugrunde. Es sollte festgestellt werden, was für eine Rolle bei kaltem Wetter aufgesuchte Schutzgebiete (Fließgewässer, Nähe zu Städten, generell wärmere Gebiete und solche mit einem höheren Feuchtgebietsanteil) sowie ihr jeweiliger Naturschutz-Status für die Artenvielfalt, die absoluten Individuenanzahlen und deren Entwicklungen auf Quartiers-Ebene spielen. Die beobachteten Zuwächse an Arten- und Individuen-Zahlen liegen auf einer Linie mit der wachsenden Bedeutung dieser Gebiete als Winterquartiere; in einigen Quartieren scheint es allerdings eine Art Dichte-abhängige Regulierung zu geben, da dort die mittleren absoluten Zahlen seit den 1990er Jahren nicht angestiegen sind. Eine Abhängigkeit von der Individuen- und Arten-Dichte spiegelt sich wahrscheinlich auch in der angestiegenen Anzahl von Fließgewässern in kälteren Gebieten wider. Dennoch sind die Faktoren, die die Entwicklung des Artenreichtums beeinflussen, auf Quartiers-Ebene weniger gut vorhersagbar. Wir zeigen die große Bedeutung von Kaltwetter-Schutzgebieten auf; für die Anzahlen an Arten und Individuen spielen Fließgewässer, wärmere Gebiete, ein höherer Anteil an Feuchtgebieten sowie die Nähe zu Städten eine große Rolle, wohingegen gesetzliche Schutzregelungen wie zum Beispiel das Ausweisen eines Naturreservats oder eines besonderen Schutzgebiets („Special Protection Area“, SPA) keinerlei Auswirkungen zeigten. Somit ist die Bedeutung von Temperatur und Kaltwetter-Schutzgebieten (Gebiete, in denen die Strenge des Winters etwas abgemildert wird) von größerer Bedeutung für die wachsende Anzahl der am Rand der traditionellen Quartiere überwinternden Wasservögel als der nachlassende Erschließungsdruck durch die Menschen.

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Acknowledgments

We are very grateful to all of the volunteers who have been involved in waterbird counts and also to the census co-ordinators of the International Waterbird Census (IWC) in the Czech Republic (i.e. to Bohuslav Urbánek, Vladimír Fiala, Čestmír Folk, Josef Křen, Ivana Kožená and Jitka Pellantová). We also extend our thanks to the Czech Society for Ornithology for its help with organizing IWC in the Czech Republic. We are grateful to Steve Ridgill for language improvement and also thank to David Storch for useful comments to earlier versions of our manuscript. This study was supported by the Ministry of Education, Youth and Sport, Project C22: “Support of PhD students and post-docs in University of Economics and Czech University of Life Sciences in Prague” and the Ministry of Environment of the Czech Republic, Project VaV MŽP ČR SP/2d3/109/07 entitled “The long-term changes in numbers and distribution of waterbirds in the Czech Republic in relation to climatic and environmental changes”.

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Correspondence to Zuzana Musilová.

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Musilová, Z., Musil, P., Zouhar, J. et al. Long-term trends, total numbers and species richness of increasing waterbird populations at sites on the edge of their winter range: cold-weather refuge sites are more important than protected sites. J Ornithol 156, 923–932 (2015). https://doi.org/10.1007/s10336-015-1223-4

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