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A Comparative Analysis of Decline in the Distribution Ranges of Orchid Species in Estonia and the United Kingdom

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A Comparative Analysis of Decline in the Distribution Ranges of Orchid Species in Estonia and the United Kingdom BIOLOGICAL CONSERVATION 129 (2006) 31– 39 available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/biocon A comparative analysis of decline in the distribution ranges of orchid species in Estonia and the United Kingdom Tiiu Kulla,*, Michael J. Hutchingsb aInstitute of Agricultural and Environmental Sciences, Estonian Agricultural University, Riia 181, Tartu 51014, Estonia bSchool of Life Sciences, University of Sussex, Falmer, Brighton, Sussex BN1 9QG, UK ARTICLE INFO ABSTRACT Article history: National databases were interrogated to analyse and compare proportional alterations in Received 14 February 2005 the distribution ranges of orchid species between two surveys in the UK (surveys com- Received in revised form pleted in 1969 and 1999) and in Estonia (surveys completed in 1970 and 2004). Nearly every 26 August 2005 species declined between the surveys in both countries, and two species may have become Accepted 21 September 2005 extinct in the UK. Mean decline in distribution range for 49 species in the UK was 50% Available online 17 November 2005 (range 14–100%), and 23 species declined by over 50%. The mean decline for 33 orchid spe- cies in Estonia was 25% (range 0–62%), and three species declined by over 50%. These Keywords: results corroborate serious range declines recently reported for orchids in other regions Distribution ranges of Europe (the Netherlands and Flanders, Belgium). In contrast with these other regions, Extinction risk we found that species associated with calcareous grassland and woodland habitats had Orchid decline suffered greater mean contractions in range than species of wet grassland habitats. Vulnerable species Greater decline was recorded for species found on drier soils, and for species characteristic Dynamic chorology of open habitats. In addition, greater decline was found in species with short inflores- cences, and in species that were short-lived, and clonal. Our results suggest that levels of decline shown both by groups of species associated with specific habitat types, and by particular species of orchid, depend strongly on local policies and specific conservation action, and indicate the habitat types on which conservation efforts may need to be con- centrated in the future. The results suggest that grazing and mowing of competing vege- tation, and avoidance of substrate disturbance, will produce the greatest rewards for the most vulnerable species. Ó 2005 Elsevier Ltd. All rights reserved. 1. Introduction increased risk of further local population decline and extinction (Fischer and Sto¨ cklin, 1997; Fischer and Matthies, The abundance and distribution ranges of many plant spe- 1998; Matthies et al., 2004). Despite considerable informa- cies have undergone dramatic declines in recent decades tion having been assembled about reductions in the ranges (Gaston, 2002; Kull et al., 2002). Important factors that have of many species, it is not clear whether decline has been contributed to these declines include habitat loss and frag- similar for groups of species characteristic of different hab- mentation (Fischer and Sto¨ cklin, 1997), and the smaller pop- itat types, or whether particular species have suffered sim- ulation sizes and greater degree of population isolation ilar levels of decline in different parts of their range. Such caused by these processes are also strongly associated with information would allow limited funds and effort to be * Corresponding author: Tel.: +372 5168512; fax: +372 7383013. E-mail address: [email protected] (T. Kull). 0006-3207/$ - see front matter Ó 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2005.09.046 32 BIOLOGICAL CONSERVATION 129 (2006) 31– 39 more closely focussed on conserving species that are at database held at the Institute of Zoology and Botany of greatest risk of extinction, both locally and over a wider the Estonian Agricultural University in Tartu. In both coun- area. tries the presence of orchid species was recorded on maps The availability of detailed records of changes in the dis- in grid squares 100 sq. km in size. Decline in distribution tribution of species over time provides opportunities for range was measured for all 49 orchid species in the UK comparative analyses of declines in ranges both within and for 33 orchid species in Estonia, and compared for and between ecological and taxonomic groupings, and be- the 30 species that were common to both countries. Decline tween geographical areas. Pioneering work in this field for each species in each country was measured as the has recently been published by Jacquemyn et al. (2005), change in the number of grid squares occupied in the na- who used records of the presence of orchid species in grid tional plant atlas databases between the two survey periods cells in Flanders, Belgium (based on 4 · 4 km cells, and re- (between 1930 and 1969, and between 1987 and 1999 in the cords in 1930 and 2000) and in the Netherlands (based on UK, and between 1921 and 1970, and between 1971 and 5 · 5 km cells, and records in 1950 and 2000) to examine de- 2004 in Estonia. Thus, there was an interval of 30 and 34 cline in their range. In Flanders, over the time period con- years between the ends of the first and second survey peri- sidered, 26 species (over 80% of those for which there ods in the UK and Estonia, respectively. The data used in were data) suffered range contractions, and eight became the analyses do not include information on the abundance extinct. In the Netherlands, 29 (78%) of the orchid species of species in each grid square. For practical purposes, in suffered range contractions, and five became extinct. Jac- large-scale comparative analyses of species decline, such quemyn et al. also compared mean percentage changes in as this, it has to be assumed that populations of all species the distribution of orchid species characteristic of different are equally detectable in all habitats (see Ke´ry, 2004 for dis- types of habitat. In both Flanders and the Netherlands, cussion of the problems associated with violation of this orchids of wet grasslands and heathlands suffered greater assumption). levels of decline than orchids of calcareous grasslands or To determine whether the magnitude of alteration in forest habitats. speciesÕ ranges was influenced by environmental prefer- The purpose of the present study was to conduct a com- ences, Ellenberg indicator values (Ellenberg et al., 1991) parative analysis of decline in the distribution ranges of spe- were tabulated for as many of the species as possible, using cies of orchids for two more widely separated European the following abiotic variables: light preference, ranging regions, namely the United Kingdom in western Europe, and from 1 (tolerant of full shade) to 9 (found in completely un- Estonia in NE Europe. The UK supports a high human popula- shaded conditions); temperature, ranging from 1 (cold-toler- tion density, and the associated infrastructure has caused ant) to 9 (tolerant of extremely warm conditions); considerable fragmentation and isolation of the remaining continentality, ranging from 1 (euoceanic) to 9 (eucontinen- patches of semi-natural vegetation, many of which appear tal); soil wetness, ranging from 1 (tolerant of very dry soils) to be suffering considerable biodiversity loss. In contrast, to 12 (plants growing beneath water); soil reaction, ranging Estonia has the lowest human population density in Europe, from 1 (highly acidic) to 9 (highly calcareous); nitrogen de- and maintains a high proportion of its semi-natural habitats mand, ranging from 1 (soils extremely nitrogen-deficient) in a protected state. Species range declines and loss of biodi- to 9 (soils extremely rich in nitrogen). In addition, the spe- versity to date in Estonia are thought to be modest compared cies were categorized according to the habitat type in which with the UK. Our aim was to compare the range declines they occur most commonly: woodlands (w), calcareous shown by individual orchid species, and groups of orchid spe- grassland (c) or wet grassland (wg). In addition, Flora Euro- cies categorised by their environmental preferences and life- paea (Tutin et al., 1980) was used to categorise species by history traits, in these two contrasting countries. Comparison mean height of inflorescence as either short (<30 cm) or tall of such widely separated regions also enabled us to ascertain (>30 cm). Information from the literature was used to cate- whether specific environmental factors and species traits gorise species as either clonal or non-clonal (whether a spe- have the same impact on decline over a wider geographical cies is capable of clonal spreading), and both published and range than that considered by Jacquemyn et al. (2005). Identi- unpublished estimates of life span were used to classify fication of environmental factors and species traits that are species as either short-lived (defined as those in which correlated with range decline may enable recommendations genet half-life is <3 years) or long-lived (those in which for management to be made that will minimise or reverse genet half-life is >3 years). The full database on which the range contraction. analysis is based is presented in Appendix 1. Statistical analysis was carried out using the SAS GLM 2. Materials and methods procedure to determine the influence of different parame- ters on decline in the range of each species. The number The area of the UK is 245,000 sq. km, and its flora includes al- of grid cells occupied at the second survey was treated as most 50 orchid species occurring in a wide range of habitat the dependent variable, with the number of grid cells occu- types. With an area of 45,000 sq. km. Estonia is only 18% as pied at the first survey as a covariate. The effects on species large as the UK, but it is rich in orchids, with 36 species re- decline of environmental factors and biotic variables, and corded nationally (Leht, 1999).
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