title: Large- and small-scale environmental factors drive distributions of cool-adapted plants in karstic microrefugia.
abstract: Dolines are small- to large-sized bowl-shaped depressions of karst surfaces. They may constitute important microrefugia, as thermal inversion often maintains cooler conditions within them. This study aimed to identify the effects of large- (macroclimate) and small-scale (slope aspect and vegetation type) environmental factors on cool-adapted plants in karst dolines of East-Central Europe. We also evaluated the potential of these dolines to be microrefugia that mitigate the effects of climate change on cool-adapted plants in both forest and grassland ecosystems.
  
  
  
  We compared surveys of plant species composition that were made between 2007 and 2015 in 21 dolines distributed across four mountain ranges (sites) in Hungary and Romania. We examined the effects of environmental factors on the distribution and number of cool-adapted plants on three scales: (1) regional (all sites); (2) within sites and; (3) within dolines. Generalized linear models and non-parametric tests were used for the analyses.
  
  
  
  Macroclimate, vegetation type and aspect were all significant predictors of the diversity of cool-adapted plants. More cool-adapted plants were recorded in the coolest site, with only few found in the warmest site. At the warmest site, the distribution of cool-adapted plants was restricted to the deepest parts of dolines. Within sites of intermediate temperature and humidity, the effect of vegetation type and aspect on the diversity of cool-adapted plants was often significant, with more taxa being found in grasslands (versus forests) and on north-facing slopes (versus south-facing slopes).
  
  
  
  There is large variation in the number and spatial distribution of cool-adapted plants in karst dolines, which is related to large- and small-scale environmental factors. Both macro- and microrefugia are therefore likely to play important roles in facilitating the persistence of cool-adapted plants under global warming.
authors.affiliation: Department of Ecology, University of Szeged, Közép fasor 52, H-6726 Szeged, Hungary zbatory@gmail.com.
authors.forename: Zoltán
authors.initials: Z
authors.lastname: Bátori
authors.affiliation: Department of Botany, Eszterházy Károly University of Applied Sciences, H-3300 Eger, Leányka utca 6, Hungary.
authors.forename: András
authors.initials: A
authors.lastname: Vojtkó
authors.affiliation: Aggtelek National Park Directorate, Tengerszem oldal 1, H-3758 Jósvafő, Hungary.
authors.forename: Tünde
authors.initials: T
authors.lastname: Farkas
authors.affiliation: Apáthy István Association, strada Baia Mare 59, RO-400171 Cluj-Napoca, Romania.
authors.forename: Anna
authors.initials: A
authors.lastname: Szabó
authors.affiliation: Milvus Group, Bird and Nature Protection Association, strada Márton Áron 9/B, RO-540058 Tîrgu Mureş, Romania.
authors.forename: Krisztina
authors.initials: K
authors.lastname: Havadtői
authors.affiliation: DRI Department of Tisza River Research, MTA Centre for Ecological Research, Bem tér 18/C, H-4026 Debrecen, Hungary.
authors.forename: Anna E
authors.initials: AE
authors.lastname: Vojtkó
authors.affiliation: Department of Ecology, University of Szeged, Közép fasor 52, H-6726 Szeged, Hungary.
authors.forename: Csaba
authors.initials: C
authors.lastname: Tölgyesi
authors.affiliation: Department of Ecology, University of Szeged, Közép fasor 52, H-6726 Szeged, Hungary.
authors.forename: Viktória
authors.initials: V
authors.lastname: Cseh
authors.affiliation: Institute of Ecology and Botany, MTA Centre for Ecological Research, Alkotmány út 2-4, H-2163 Vácrátót, Hungary.
authors.forename: László
authors.initials: L
authors.lastname: Erdős
authors.affiliation: Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza Street 64, 00-679 Warsaw, Poland.
authors.forename: István Elek
authors.initials: IE
authors.lastname: Maák
authors.affiliation: School of Natural and Built Environments, University of South Australia, Mawson Lakes Campus, GPO Box 2471, Adelaide, South Australia 5001, Australia.
authors.forename: Gunnar
authors.initials: G
authors.lastname: Keppel
mesh_terms: D057231:Climate Change; D003080:Cold Temperature; D017753:Ecosystem; D018521:Plant Physiological Phenomena; D000068236:Refugium
keywords: Capacity of refugia; East-Central Europe; environmental gradient; high-mountain plants; karst dolines; migration processes; refugia; relicts; slope aspect