Acta Oecologica 36 (2010) 548e560

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Acta Oecologica
journal homepage: www.elsevier.com/locate/actoec

Original article

Relevance of multiple spatial scales in habitat models: A case study with amphibians and grasshoppers
Michael Altmoos*, Klaus Henle
UFZ e Helmholtz-Centre for Environmental Research, Department of Conservation Biology, Permoserstrasse 15, D-04318 Leipzig, Germany

a r t i c l e i n f o
Article history: Received 8 April 2009 Accepted 11 August 2010 Available online 9 September 2010 Keywords: Amphibia Conservation planning Habitat model Habitat suitability Orthoptera Spatial scales

a b s t r a c t
Habitat models for animal species are important tools in conservation planning. We assessed the need to consider several scales in a case study for three amphibian and two grasshopper species in the postmining landscapes near Leipzig (Germany). The two species groups were selected because habitat analyses for grasshoppers are usually conducted on one scale only whereas amphibians are thought to depend on more than one spatial scale. First, we analysed how the preference to single habitat variables changed across nested scales. Most environmental variables were only signicant for a habitat model on one or two scales, with the smallest scale being particularly important. On larger scales, other variables became signicant, which cannot be recognized on lower scales. Similar preferences across scales occurred in only 13 out of 79 cases and in 3 out of 79 cases the preference and avoidance for the same variable were even reversed among scales. Second, we developed habitat models by using a logistic regression on every scale and for all combinations of scales and analysed how the quality of habitat models changed with the scales considered. To achieve a sufcient accuracy of the habitat models with a minimum number of variables, at least two scales were required for all species except for Bufo viridis, for which a single scale, the microscale, was sufcient. Only for the European tree frog (Hyla arborea), at least three scales were required. The results indicate that the quality of habitat models increases with the number of surveyed variables and with the number of scales, but costs increase too. Searching for simplications in multi-scaled habitat models, we suggest that 2 or 3 scales should be a suitable trade-off, when attempting to dene a suitable microscale. 2010 Elsevier Masson SAS. All rights reserved.

1. Introduction Habitat models for animal species are important tools for conservation planning. They are statistical models that relate the abundance or presence/absence of an animal species to environmental parameters. Model-based evaluations of habitat suitability allow those areas and structures to be identied that are of particular importance for the survival of target species (Wiegand et al., 1999; Altmoos and Henle, 2007). Habitat models are also important tools for solving conservation goal conicts that may exist for specic sites (Altmoos, 1999a, b; Drechsler, 2004). Knowledge of successional changes in habitats enables one to

* Corresponding author at: Nahe der Natur e Museum fr Naturschutz, Schulstrasse 47, D-55568 Staudernheim (Nahe), Germany. Tel.: 49 6751 8576370; fax: 49 6751 8576346. E-mail addresses: michael.altmoos@nahe-natur.net (M. Altmoos), klaus.henle@ ufz.de (K. Henle). 1146-609X/$ e see front matter 2010 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.actao.2010.08.001

predict the temporal changes in the suitability of a site for target species, hence allowing different management scenarios to be evaluated (Kliskey et al., 1999). Recently, habitat models have also been frequently used to predict impacts of climate change on species distribution (e.g., Arajo et al., 2006). The rst simple habitat models were developed by the U.S. Fish and Wildlife Service (1981) and were followed by many variations and extensions (Kliskey et al., 1999; Kleyer et al., 2000; Carter et al., 2006). They range from simple estimates to complicated computer simulations (neuronal networks, fuzzy logic) that emerged after several years of development. Most habitat models are based on univariate or multivariate statistical analyses on a single scale (Pearsall et al., 1986; Pavignano et al., 1990; Halley et al., 1996; Kliskey et al., 1999; Osborne et al., 2001). Univariate approaches enable us to understand whether particular factors inuence the occurrence of a species in the same or in different ways across scales, whereas multivariate analyses allow evaluating whether and how different scales should be combined optimally to accurately predict the presence or absence of a species.

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Spatial extrapolation of habitat models is greatly facilitated if key variables can be easily measured, ideal variables are structures that can be identied by remote sensing (Osborne et al., 2001). In such cases habitat models can greatly facilitate site evaluations (Pearce et al., 2001) and the development of effective networks of conservation priority sites (Altmoos, 1999a; Margules and Pressey, 2000). The assessment of habitat suitability conducted at just one spatial scale may be of limited use because different environmental variables may inuence habitat use and tness at different scales (Luck, 2002; Scott et al., 2002). More insights can be gained by investigating habitat suitability at multiple spatial scales (Hall and Mannan, 1999; Storch, 2002; Pellet et al., 2004; Schrder et al., 2007). The multi-scale approach acknowledges the inuence of spatial variation on species, recognising that the answers we get depend on the scale we use to answer a question. This means that using one scale only may not be wrong, but it may not provide enough explanatory power and may only partially answer a question (Morris, 1987; Levin, 1992; Otis, 1997; Melbourne et al., 2004). For example, understanding metapopulation structure and dispersal in heterogeneous landscapes (Hanski and Gaggiotti, 2004) requires us to assess the suitability of habitats from the scale of an individual home range to the landscape scale at which dispersal takes place (examples for amphibians: Fog, 1993; Hartel et al., 2008). Such multi-scale considerations are particularly important in highly dynamic landscapes (Kleyer et al., 2007). Although multi-scaling is important, it can complicate the collection and analysis of data because more variables may have to be surveyed. It is often easier to measure and analyse habitat variables on one scale only and this may explain why habitat models do not yet routinely include several scales. If multi-scale habitat models are to be broadly used in management, a feasible trade-off must be found between a sufcient reection of natural spatial complexity and schematic simplication (Karl et al., 2000; Pearce et al., 2001). Few studies have systematically assessed trade-offs or compared the relevance of multi-scaling among different taxonomic groups (Scott et al., 2002). Grant et al. (2004) showed that one scale may be enough for butteries, but not for birds. Gibson et al. (2004) concluded from a habitat study of rufous bristlebirds (Dasyornis broadbenti) in Australia that habitat models with more than one scale would be much better for conservation planning, even when a habitat model on one particular scale is already used and suitable for the rst steps in practical conservation. Searching for generally feasible simplications in conservation planning, we used species from two very different taxonomic groups (amphibians and grasshoppers) to test a) whether habitat variables are equally suitable across scales, whenever they can be measured at more than one scale, b) whether variable combinations within a single scale provide sufcient accuracy for predictions, c) whether the combination of variables at different scales enables more accurate predictions with the same amount of variables or less than on a single scale, d) whether for the selected taxa, a multi-scaling approach is really necessary to obtain suitable minimum models, i.e., models that use as few variables as possible.

a single scale (Kuhn and Kleyer, 2000), whereas for amphibians habitat suitability is usually considered both at the pond and the landscape scale (Gnther, 1996). Besides this, their dispersal distances, foraging ecology, and sexual behavior indicate that analyzing more than one scale may be useful. We limited our study to species that can be reliably detected and recorded using standardized audio and visual surveys (Ingrisch and Khler, 1998; for a detailed description of the detection methods for each species see Altmoos, 1999b):  The European tree frog (Hyla arborea L., 1758.), the green toad (Bufo viridis Laurenti, 1768), and the spadefoot toad (Pelobates fuscus Laurenti, 1768) are under threat across large parts of their German distribution, but occur commonly in the study area. They are already known as multi-scaling organisms (Gnther, 1996; for the European tree frog: Pellet et al., 2004). We limited our analysis to potential breeding habitats and used calling males as an indication of breeding.  The blue-winged grasshopper Sphingonotus caerulans (L., 1767) and Oedipoda caerulescens (L., 1758) are ground-dwelling species in open and dry grassland. Past habitat analyses for these species were made on the basis of one spatial scale only (e.g., Kuhn and Kleyer, 2000).

2.2. Study region Our study region is an approximately 200 km2 opencast lignite-mining landscape situated in the 30 km 30 km mining region south of Leipzig (Central Germany). This landscape contains 128 mining sites (for a detailed description of all sites see Altmoos, 1999b). We selected non-recultivated opencast mining landscapes as a model system because they provide complex dynamic habitats for many endangered species (Altmoos and Durka, 1998), each with its own specic habitat requirements. We used four large sites from the 128 opencast mining sites (Bockwitz, Lobstdter Lachen, Werben-Sittel pit, and the North Profen dump) to develop habitat models. 2.3. Spatial scale approach We distinguished three spatial scales each for amphibians and grasshoppers (microscale, mesoscale, and macroscale: Fig. 1). Our approach is similar to that of Luck (2002) and Caruso et al. (2005) but adjusted to our study organisms and landscape. The three scales were as follows for amphibian calling sites (listed from smallest to largest):  Microscale is the place with a radius of 5 m around a site where males called. Such small scales have rarely been studied for amphibians. We selected a radius of 5 m because at larger scales the position of calling males is probably inuenced by the location of competing males and because at smaller scales some habitat structures that are assumed to be important for these species cannot be quantied.  Mesoscale: entire pond, the size of which may vary; entire ponds traditionally represent the scale, which has been investigated most intensively for amphibians (e.g., Laan and Verboom, 1990; Pavignano et al., 1990). The smallest ponds were approx. 5 m2, the largest pond was approx. 10,000 m2, the typical ponds (90% of all water bodies) were between approx. 10 and 200 m2.  Macroscale refers to the area that surrounds a pond. This scale features large-scale landscape variables, which are generally regarded as important for amphibians because of

2. Materials and methods 2.1. Model species We chose grasshoppers and amphibians as model taxa, because the habitat requirements of grasshoppers are usually assessed on

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Fig. 1. Levels of nesting of the spatial scales used in the habitat analyses.

their summer home ranges (Osborne et al., 2001; Pellet et al., 2004). We used two buffers around the pond, one of 300 m and one of 3 km. We chose these buffers because they cover the main dispersal distances of amphibians from breeding sites (Fog, 1993; Gnther, 1996) and should therefore represent a suitable macroscale. The spatial scales used for grasshoppers are as follows:  Microscale of 1 m2 quadrants e the smallest spatial scale at which habitat variables can be quantied adequately and which may be relevant for the physiology of grasshoppers (Appelt, 1996; Ingrisch and Khler, 1998).  Mesoscale of 25 m2 grids e often reecting the area actively used by grasshoppers (Appelt, 1996; Samietz and Berger, 1997).  Macroscale of 5000 m2 area e often used in habitat descriptions for grasshoppers to include dispersal abilities and landscape inuences (Ingrisch and Khler, 1998). For surveying amphibians, we placed transects through the four mining sites that covered all 199 ponds within these sites. Each

pond was surveyed 10 times to ensure that every pond was surveyed at an optimal time. At each pond, and every 100 m for large ponds, we stopped for 10 min to locate calling males. We also used the same procedure to survey unoccupied sites e only those sites that were at least 10 m away from a calling male were considered to be unoccupied. We stopped selecting unoccupied sites once we had located 400 such sites. At each of these sampling points, we measured habitat variables at all three scales (Fig. 1). This approach showed that 97 of the 199 ponds were used for breeding by at least one of our species. In total, we obtained 188 calling points for B. viridis, 513 for P. fuscus, and 213 for H. arborea (see Appendix). For surveying grasshoppers, we also placed transects through open areas across all four mining sites. The starting points and the directions of transects were randomly determined for each survey. We visually surveyed grasshoppers along the transects until we obtained 1000 occupied points for each species. We repeated the same procedure to survey 1000 points that were not occupied by any of our grasshopper target species. We determined habitat variables at each selected point across all three scales (Fig. 1).

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In presence/absence, surveys a low detection probability may create false absences (Pellet et al., 2004) and this could inuence the precision of habitat models (MacKenzie et al., 2006). We therefore conducted surveys under optimal conditions, made extensive recordings at each sampling point, the search area was small, and there was a high visibility at our study sites. Therefore, the detection probability should be very close to 1. The high prediction accuracy obtained from our nal habitat models supports this conclusion. 2.4. Habitat variables at different scales Based on literature (Nllert, 1990; Gnther, 1996; Ingrisch and Khler, 1998; Khler, 1999), we selected 19 potentially relevant habitat variables for amphibians (Table 1) and 11 variables for grasshoppers (Table 2). We decided to make as few restrictions as possible at this stage in order to avoid accidentally missing certain variables, which may prove to be important for habitat modelling. However, we only included those factors, which can be surveyed easily, since this is of paramount importance for applied conservation. On each scale, we measured all the variables in the eld using standard survey techniques (see Appendix for overview; for a detailed description see Altmoos, 1999b): We visually classied the vegetation cover, bare ground and degree of shade into bands of 10%. The cover is related to the whole area at the scale analysed. We averaged the abiotic water characteristics across sampling points spaced 10 m apart along the entire edge of the water. 2.5. Habitat models To explore the relevance of scales for habitat models, we used both univariate and multivariate approaches. Both approaches are common in habitat models (e.g., in determining habitat suitability indices: U.S. Fish and Wildlife Service, 1981; Pearsall et al., 1986), but they convey different information that is relevant for applied conservation and for understanding the habitat requirements of species. Univariate approaches clearly show the preference for single factors that had been recognized as important, particularly in homogenous habitats. They are easier to translate into habitat suitability indices that are easy to understand and apply for practitioners (Pearsall et al., 1986). Multivariate methods, in contrast, enable a much better description of the preferences in more complex habitats, in which different habitat variables may have interacting effects. 2.5.1. Univariate preference analysis We used Spearmans rank correlation tests pair-wise among all factors and on each scale. This would enable us to lter out replaceable factors from further consideration. Replaceable factors are variables that are highly correlated with other variables. For amphibians, only the degree of cover of submerged and emergent vegetation showed a signicant correlation (pond scale; r 0.84). Since these two factors relate to different requirements for amphibians (submerged vegetation as a substrate for depositing eggs; emergent vegetation to provide cover and calling sites for H. arborea), we retained both factors for further analyses. We examined preference or avoidance for each habitat variable on each scale. We compared the number of occupied places for the categories of a habitat variable with the expected number of occupied places given the prevalence of the habitat categories among all analysed sites using a c2-test.

Since we carried out a total of 111 univariate tests for amphibians and 50 for grasshoppers, we used a signicance level of a < 0.01 for the univariate tests. Therefore, we expect approximately one or two results to be signicant by chance. Alternatively, we could have used a Bonferroni adjustment procedure, but we preferred the former approach because it provides a better balance between type I and type II errors. 2.5.2. Multivariate modelling techniques and optimal habitat models In order to investigate not just the inuence of individual factors but entire scales, we produced multivariate logistic regression models (Hosmer and Lemeshow, 1995) separately for each spatial scale using all variables measured at that scale as well as for all combinations of scales. We did not include sites as a xed factor nor did we incorporate autocorrelation into our models because we wanted to keep our models as simple as possible. Side-identity and spatial autocorrelation are proxies for unmeasured process variables that determine the distribution of species. Our models already provided very high prediction accuracy for most species (see Results) without the inclusion of site identity or autocorrelation parameters. Thus, unmeasured process variables reected in the spatial structure of the data could only explain additional variability to a limited extent. Most likely, this is due to the high diversity within mining sites and similarity among mining sites. Moreover, spatial regression methods tend to emphasize smallscale patterns whereas non-spatial regression methods emphasize broad-scale patterns (Diniz-Filho et al., 2007) but our non-spatial methods already demonstrate the importance of the microscale (see Results) so that the use of spatial models should strengthen our conclusions. Note that parameter estimates obtained with non-spatial models are unbiased and only error terms are affected, which are not relevant in our study because our models were not used for hypothesis testing. In the next step using backward selection, we deleted as many variables as possible from the model that included all variables at all scales to obtain a minimum or optimal model that contained as few variables as possible but still achieved a sufcient predictive accuracy. Such models can be regarded as an optimal stage for use in conservation planning because they reduce the amount of material and costs in eld surveys for similar landscapes. Variables that did not improve the model signicantly (a < 0.05) were removed. We rst removed the variable, which contributed least to the explanation of the model and then the variable that contributed least to the resulting model. Following Hosmer and Lemeshow (1995) and Carter et al. (2006), we stopped removing variables when the resulting model predicted less than 85% of the presences and absences correctly; the percentage of correct predictions is the correct classication rate (CCR). We used SPSS for performing the multivariate logistic analyses. Our criteria are stringent and favour models with only a few variables. With such stringent criteria no logistic regression model may be acceptable. This was the case for H. arborea and O. caerulescens. Rather than relaxing the stringency of the criteria set for these species, we followed the simple approach used by the U.S. Fish and Wildlife Service (1981) for applied habitat models by translating the statistically signicant results from the univariate analyses into a habitat suitability index between 0 and 1. In terms of modelling, this standardisation naturally represents fuzzy estimates, which must not be interpreted as mean values. To link up the variables, we added up the suitability indices to get an overall index as suggested by Pearsall et al. (1986). We added the variables stepwise testing all possible combinations until the prediction accuracy reached at least 85%.

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Table 1 Summary of statistically signicant preferences and avoidances of habitat variables for amphibians. Variables Species Case no. Microscale (5 m radius) Preferences pH BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF BV HA PF 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 e e e e e e e e e e e e e e e e e e High High e 0.1e0.5 m 0.1e0.5 m e >80% e <10% e e e e e e <20% e e e e e 0e20% 20e40% 20e40% 0e20% 20e60% 20e60% e e e Low Low e >0.5 m >0.5 m e e e e e e e e e e e e e >80% e e >40% >60% 60e80% >20% <20%, >80% <20%, >80% e e e >6 >6 >6 e e e e e e Avoidances <6 <6 <6 e e e e e e Mesoscale (pond) Preferences >6 >6 >6 e e e e e e No sh No sh No sh 3e10 years e e e High e e 0.1e0.5 m e e e >10%, <80% e e e e e e e e e e e e 20e40% 20e60% 60e80% e 20e40% e e e e e e e e e e Avoidances <6 <6 <6 e e e e e e e e e e e e e Low e >0.5 m >0.5 m e e e e e e e e e e e e e <40% <40% e >60% e <20% e <20% <20%, >60% e e e e e e e e e e Forest closer 3 km e e >5 Closer 3 km e e <300 m e e e e e e e e e e e e e e e e Macroscale (landscape) Preferences Avoidances

Conductivity

Nitrogen, total

Presence of sh

Age

Warming capacity

Maximum depth

Fluctuation of water level

Turbidity

Water ow

Slope of bank (inclination > 20 are steep banks) Maximum degree of shading

Coverage of submerged vegetation

Coverage of emergent vegetation

Composition of soil types

Pond size

Habitat types in surrounding landscape

Number of ponds in area (<300 m and 3 km radius) Distance to next pond

BV Bufo viridis; HA Hyla arborea; PF Pelobates fuscus. The survey scheme, the numbers of sites, and the tests are explained in the text (see Materials and methods). blank Variable not dened or measured. d Variable measured, but no signicant preference or avoidance (variable is not predictive).

M. Altmoos, K. Henle / Acta Oecologica 36 (2010) 548e560 Table 2 Summary of statistically signicant preferences and avoidances of habitat variables for grasshoppers. Variables Species Case no. Microscale (1 m2) Preferences Total vegetation coverage Vegetation coverage 5 cm above ground Vegetation coverage 20 cm above ground Vegetation coverage 50 cm above ground Percentage of tree shading Percentage of bare ground Soil type Slope Exposition (if slope > 20 ) Orb spider abundance class Age OC SC OC SC OC SC OC SC OC SC OC SC OC SC OC SC OC SC OC SC OC SC 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 e e e e e e e e 10e20% <10% 10e40% 10e20% <10% <10% e <10% <10% 0% Avoidances >60% >40% >40% >30% >30% >20% >20% >10% >10% >0% Mesoscale (25 m2) Preferences 20e50% 10e40% 30e70% e e <10% e e 1e30% 1e30% >80% 50e70% e e e e e e e e e 3e10 Years Avoidances >70% >60% >80% e e >10% e >10% >50% >20% <20% e e e e e e e >3 >3 e e e 3e10 Years e e e e e e e e e e e e >60% >20% e e e e e e e e Macroscale (5000 m2) Preferences 30e70% 20e40%

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Avoidances <20% >60%, <20%

OC Oedipoda caerulescens; SC Sphingonotus caerulans. The survey scheme, the numbers of sites and the tests are explained in the text (see Materials and methods). blank Variable not dened or measured. e Variable measured, but no signicant preference or avoidance (variable is not predictive).

3. Results 3.1. Univariate preference analysis Thirteen of the 19 variables measured for amphibians and 8 of the 11 variables measured for grasshoppers were signicantly preferred or avoided (c2-tests; a < 0.01) by at least one species at one scale. The combination of habitat variables and species resulted in 57 cases for amphibians and 22 cases for grasshoppers, in which preferences can be analysed across scales (Tables 1 and 2). In 25 of the 57 cases for amphibians and in 15 of the 22 cases for grasshoppers, we observed a signicant preference or avoidance on at least one scale. In 21 cases for amphibians and in ten cases for grasshoppers, variables were predictive on at least two scales. Preferences or avoidances of the habitat variables differed among scales. We classied species-specic effects into four categories (Fig. 2): (a) Similar multi-scale preferences. In no case did preferences remain exactly constant across scales but in 13 cases (eight amphibian cases, Table 1, and ve grasshopper cases, Table 2), variables were equally relevant across scales: only the peaks of the preferences shifted along the variable axis as illustrated for O. caerulescens in Fig. 2A. It preferred low total vegetation cover across all scales; at the microscale, the peak of preference was at 20e30%, at the mesoscale 30e40%, and at the macroscale 40e60% total vegetation cover. (b) Singularities e the occurrence of preferences only on a particular scale and new variables per spatial scale. In 12 cases for amphibians and in two cases for grasshoppers, the

variables were only measured at one scale (Tables 1 and 2). For example, factors of distance and landscape variables can of course only be measured at the macroscale and were decisive for H. arborea. The age of a pond is also by denition only measurable at the level of the entire pond, with B. viridis preferring ponds aged 3e10 years old. The grasshopper S. caerulans preferred 51e70% bare ground (51e70%) at the mesoscale (Fig. 2B) but the variable was not predictive at the macroscale. (c) Signicant differences for preferences at different scales. A signicant preference at one spatial scale that is no longer significant at another scale occurred in nine cases for amphibians (Table 1) and in eight cases for grasshoppers (Table 2). The difference between the two groups is marginally signicant (Fishers exact probability: p 0.08). For example, B. viridis showed a preference for water with high warming capacity, a water depth of 0.1e0.5 m, high uctuations of the water level, and places without a steep bank at the microscale but not at the mesoscale. It also avoided submerged vegetation cover above 1e20% at the microscale but not at the mesoscale, where it preferred a submerged vegetation cover of approx. 20e40% (Fig. 2C). (d) The reversed effect of individual variables at different spatial scales. In one amphibian and two grasshopper cases, shifts involved the reversal of preference and avoidance (case no. 39 in Table 1 and cases no. 3 and 10 in Table 2). The difference between grasshoppers and amphibians is signicant (Fishers exact probability: p 0.02). For example, P. fuscus preferred a high cover of submerged vegetation of 60e80% at the mesoscale but signicantly avoided this vegetation cover at the microscale; instead, at this scale, the species preferred

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Fig. 2. Classication of univariate habitat preferences across scales. The higher the peak the more suitable is the habitat. Preferences: as a measure of preference or avoidance we used the difference between observed presence (occupation of species) and the expected presence (the presence of the feature in the reference data set) at each class [%]. Statistical explanations are in the method chapter.

submerged vegetation cover of 20e40% (Fig. 2D). Therefore it can be said that P. fuscus prefers well-vegetated ponds as breeding sites (reproduction), and yet for calling they select places with sparse vegetation (compare Nllert, 1990). Both, low and high vegetation cover at the same pond, but at different spatial scales, are necessary and this can only be recognized with the multi-scale approach.

3.2. Multivariate models: combinations of scales and multi-scale effects on habitat models The highest possible classication accuracy is of course achieved by incorporating all habitat variables at all scales (Table 3). In these models, the correct classication of occupied sites was

92.9% for S. caerulans, 92.3% for P. fuscus, and 91.3% for B. viridis. By contrast, for H. arborea and O. caerulescens only 65% respectively 82.2% was achieved, which is not sufcient under our stringent criteria. Hence, no suitable model could be derived by using multivariate logistic regressions for these two species. H. arborea proved to be a unique case in which only occupied places could not be predicted with sufcient correctness. Unsuitable places could be identied relatively accurately with a rate of 92.1%. Information lost from disregarding spatial scales increased irregularly as the number of spatial scales that were disregarded increased and varied sharply among species (Table 3). For example, when only one spatial scale was considered for H. arborea, only a 53.3% correct classication was achieved with all variables at the microscale and 45.9% with all variables at the

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Table 3 Maximum correct classication rates (CCR) for occupied and non-occupied (in parenthesis) sites for various combinations of scales. On every scale all variables were considered in multivariate logistic habitat models. Grey shading: models with a CCR > 85%; n: sample size. Species Single scales (scale numbera) 1 Hyla arborea (n 613: 400 empty 213 occupied places) Pelobates fuscus (n 913: 400 empty 513 occupied places) Bufo viridis (n 588: 400 empty 188 occupied places) Sphingonotus caerulans (n 2000: 1000 empty 1000 occupied places) Oedipoda caerulescens (n 2000: 1000 empty 1000 occupied places)
a

Combination of two scales (scale numbersa) 12 (28.0) (12.4) (22.3) (49.2) (56.7) 58.8 91.0 90.0 91.1 78.1 (65.0) (85.1) (93.7) (87.0) (75.6) 13 60.8 89.2 89.3 82.2 72.7 (80.2) (82.0) (87.2) (78.8) (70.0) 23 55.6 79.0 85.4 88.4 76.7 (73.5) (70.9) (88.8) (86.1) (74.4)

All scalesa 123 65.0 92.3 91.3 92.9 82.2 (92.1) (85.8) (98.9) (88.7) (76.4)

2 (70.2) (78.2) (92.7) (70.5) (53.2) 45.9 75.6 73.1 78.2 75.3 (55.5) (65.8) (77.6) (75.9) (74.1)

3 23.1 10.2 25.2 51.3 61.8

53.3 87.3 87.2 68.5 58.6

Scale numbers refer to microscale (1), mesoscale (2), and macroscale (3).

mesoscale. In contrast, the microscale alone was sufcient for correct classications for B. viridis, although it was slightly less accurate compared to when all scales were included. This made B. viridis the only one of our species, for which just one scale was sufcient to create a suitable habitat model. For two species, different combinations of two scales provided a sufcient degree of correct classications. For P. fuscus, the combination of the two lower scales achieved a 91% correct classication, which was only marginally lower than for three scales. However, a sufciently accurate model could not be derived from any other combination of two scales. Likewise, the combination of the two lower scales (micro- and mesoscale) leads to a correct presence classication of 91.1% in the case of S. caerulans. Combining the mesoscale with the macroscale also delivered a sufciently accurate model, albeit with a rating of 88.4% that is not quite as reliable as the combination of the two lower scales. In summary, different combinations of scales lead to adequate models. The dependency on individual spatial scales for a habitat model meeting the minimum quality threshold varied among species. As far as amphibians were concerned, B. viridis only required one scale for an adequate model, whereas P. fuscus required two scales, and for H. arborea not even all three scales were adequate for the set of habitat variables tested. The only general conclusions that could be drawn were that the more spatial scales that were considered, the higher the quality of the model, and that the lowest scale was particularly

important for all species. Only in the case of B. viridis did the loss of the lowest spatial scale result in a model that still offered a correct classication, which was just acceptable (85.4%) but even the quality considering the microscale alone was not achieved (Table 3). Hence, the lowest scale (microscale) was very important for the habitat models of all the species investigated here, and in fact decisive for P. fuscus. 3.3. Optimal models and different species-specic requirements of multiple scales Key variables of minimum or optimal models, i.e. models, which achieve at least 85% correct classication with as few variables as possible (irrespective of the scales used), are listed in Table 4. For H. arborea and O. caerulescens, where logistic regressions did not result in a sufcient model, we were able to nd additive models obtained from univariate preferences as outlined in the methods chapter (i.e., by transforming preferences into habitat suitability indices between 0 and 1 and by combining the variables). A single spatial scale, namely the microscale, was sufcient for minimum models only in the case of B. viridis. Four habitat variables were sufcient at this scale to provide a simple yet functional model. In the other species investigated, variables from at least two scales had to be combined. In the case of P. fuscus and S. caerulans these were the two lower spatial scales,

Table 4 Habitat variables and number of spatial scales required for optimal habitat models. Species and model Hyla arborea (n 613: 400 empty 213 occupied places) Additive connection of variables CCR 86.6 (85.4) Habitat variables [scale number]  Water depth, pH, emergent vegetation cover [#1]  Emergent vegetation cover [#2]  Number of ponds in a 300 m buffer [#3]  Distance to other ponds < 3 km [#3: larger surrounding]  pH [#1]  Submerged vegetation cover [#1]  Fluctuation of water level [#2]     pH [#1] Fluctuation of water level [#1] Shading [#1] Submerged vegetation cover [#1] Number of spatial scales required 3 (# 1 2 3)

Pelobates fuscus (n 913: 400 empty 513 occupied places) Logistic regression CCR 88.5 (85.1) Bufo viridis (n 588: 400 empty 188 occupied places) Logistic regression CCR 87.0 (96.5) Sphingonotus caerulans (n 2000: 1000 empty 1000 occupied places) Logistic regression CCR 92.0 (89.0) Oedipoda caerulescens (n 2000: 1000 empty 1000 occupied places) Additive connection of variables CCR 92.2 (88.9)

2 Scales (# 1 2)

1 Scale (# 1)

 Total vegetation cover [#1]  Coverage of bare ground [#2]  Vegetation cover at 5 cm height [#1]  Total vegetation cover [#3]

2 Scales (# 1 2)

2 Scales (# 1 3)

CCR: correct classication rate (%) of occupied and, in parenthesis, of unoccupied sites; n: sample size; # scale number (1: microscale, 2: mesoscale, 3: macroscale).

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whereas with O. caerulescens a variable at the microscale had to be combined with a variable at the macroscale. In the case of H. arborea, variables from all scales were required. 4. Discussion 4.1. Changing relevance of habitat variables with scales Similar preferences across scales will facilitate applied conservation and are often taken for granted without being tested (overview on amphibians in Germany: Gnther, 1996; grasshoppers in Germany: Ingrisch and Khler, 1998). Our univariate analyses showed that this assumption was met more as an exception than as a rule: only a few variables were equally relevant at all spatial scales, and only very rarely is one of their prevalences preferred at all scales. Preferences shifted with spatial scales in approx. 20% of cases for both taxonomic groups. Moreover, reversals between preference and avoidance across scales occurred in approx. 5% of the cases. Grasshoppers and amphibians did not differ signicantly in terms of the percentage of habitat variables that were relevant on all measured scales or in terms of the percentage of shifts with scale. From the shifts observed, two of the variables included were relevant for optimal habitat models (submerged vegetation cover for B. viridis; uctuation of the water level for P. fuscus). All three reversals observed involved variables that were relevant for optimal habitat models (submerged vegetation for P. fuscus; vegetation cover 5 cm above the ground for O. caerulescens; total vegetation cover for S. caerulans). Therefore, widespread generalised textbook statements, such as species requires vegetation cover y, must be associated with particular scales in order to avoid inaccuracies or even incorrect information regarding habitat suitability. 4.2. Why are single scale habitat models often not sufcient? While for various taxa, models are published that are based on single scales and reach sufcient predictive power, a rigorous assessment under the conditions of which a single scale may be sufcient is lacking. Notwithstanding, it is known that multiple spatial scales are essential for adequate extrapolations from habitat models in heterogeneous environments (Scott et al., 2002; Gibson et al., 2004). Our results indicate that multiple spatial scales are usually also essential for habitat models in supercially more homogeneous habitats and for species that may be supercially regarded as mono-scale species, such as grasshoppers in dry grasslands. This shows that both large-scale and small-scale habitat features determine the presence of the species being examined by us. B. viridis was an exception: For this species, a single scale was sufcient and achieved almost the same predictiveness as all three scales combined, although on each scale all potentially suitable variables were measured. Likewise on another level, for the purpose of identifying species hotspots, a single scale was sufcient for moths but not for butteries in a pitch pine-scrub oak landscape (Grant et al., 2004). Published habitat models for amphibians ponds are often based on uniscale analyses; they can reach acceptable predictive accuracy (>85% correct classications of species presence) but require many habitat variables (e.g., Pavignano et al., 1990; Halley et al., 1996). However, single scale models may also fail. H. arborea is one such example. Pellet and Schmidt (2005), who focused on a single scale, could not nd any adequate habitat model. Similarly, Van Buskirk (2005) examined six variables at the pond level and two at the landscape level and came to the conclusion that only the pond level

scale is important but obtained unsatisfactory low explanatory power. In contrast, Pellet et al. (2004) obtained higher explanatory power when testing for the necessity of a landscape scale for the same region in Switzerland and concluded that at least two scales are required. Even when using two scales with variables that are predictive there is room for improvement of the amount of variability explained, as is well demonstrated by our results for H. arborea. We conclude that different scales are often important for amphibians because without multiple scales important factors could easily be neglected. However, the relevance of different scales may depend on the objectives of modelling, the variables assessed, and the variability within the study region. Similar to our study, Luck (2002) showed for the Australian bird species Climacteris rufa that habitat preferences can only be understood when different scales are considered even if many variables could only be measured on one scale as was the case in our study. Likewise, both neighbouring water bodies within 50 m and certain plant species in the breeding pond are crucial for the occurrence of the Australian golden bell frog (Litoria aurea) (Hamer et al., 2002). Hence, multi-scaling solves the problem that is frequently encountered in the analysis of habitats when determining the relevant grid size of habitat preferences. The fact that most variables only showed signicant preferences at certain scales supports previous insights that many ecological patterns (e.g., body size abundance relationships: Blackburn and Gaston, 1997; buttery species diversity: Debinsky et al., 2001; relationship between insect density and habitat conguration: Krawchuk and Taylor, 2003) and processes (e.g., decomposition: Saunders et al., 2002; life-history traits as predictors of species sensitivity to fragmentation: Henle et al., 2004) need to be regarded from a multi-scale perspective (see also the review by Levin, 1992; Scott et al., 2002). Our results regarding the relevance of multi-scaling in speciesehabitat relationships also conform to the results of studies using analogous multi-scale approaches (Hall and Mannan, 1999; Storch, 2002; Hartel et al., 2008). The importance of different scales for habitat models can be explained with the mosaic cycle theory and metapopulation theory. According to the mosaic cycle theory, species survive in heterogeneous landscapes by tracing different successional stages of vegetation. The stages are distributed like a mosaic within the landscape and shift spatially with time (Kleyer et al., 2007). From metapopulation theory and empirical studies, it is well-known that the viability of metapopulations depend on both the population size of a patch and the spatio-temporal distribution of patches (Hanski and Gaggiotti, 2004). The population size of a patch is a function of patch habitat quality, and habitat factors on the microscale and/or the mesoscale may be decisive (this study; examples cited above). Therefore, the habitat requirements of species can be adequately described and their presence or abundance only predicted by using different spatial and temporal scales. Since it is assumed that the majority of species occur as metapopulations (Thomas et al., 2001), it is understandable, why it would be advisable to pursue multi-scale approaches to habitat models. 4.3. Species-specic importance of macro- and microscales and optimal models Since amphibian species are generally regarded as being dependent on factors surrounding ponds (our macroscale, cf. Gnther, 1996), it is somewhat surprising that this only had to be reected in the habitat models for H. arborea, whereas for the other species simplications without the macroscale were

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sufcient for correct predictions. The dependence of H. arborea on a macroscale in combination with a pond scale is already known (Pellet et al., 2004). Moreover, in all three amphibian species, the models based only on the macroscale gave the worst predictions and were completely unacceptable. A macroscale may be necessary for modelling some species like H. arborea, or for the common toad (B. bufo) in a case study in Romania (Hartel et al., 2008), but a macroscale alone is not sufcient - even for them. The lack of response to the current landscape at the 3 km level in B. viridis and P. fuscus may be explained by their good dispersal ability (Gnther, 1996), their preference for open habitats, which are characteristic for mining sites, and the higher within mining-site heterogeneity compared to among miningsite heterogeneity in our study region. For the grasshopper species, no scale dominated as clearly as for the amphibian species, although the mesoscale provided a slightly better prediction accuracy than the other two scales. Notwithstanding, only the microscale was included in optimal habitat models for both species. Hence, this scale harbours important preferences that cannot be measured as well at other scales. In contrast to our approach, conservation practice includes many habitat models, which do without microscales and hence offer less reliable predictions (e.g., Kuhn and Kleyer, 2000). As a further example, Thomas et al. (2001) showed for several buttery species that within-patch habitat quality better explains patch occupancy and abundance than patch size. This also suggests that the microscale is of particular importance. The lower the scale, the higher the natural dynamics of species presence. For example, the microscale is essentially the place where a male frog calls (calling points, see Materials and methods). These calling sites are of course not static. A change in calling sites may take place within and among seasons, but it will not inuence the general importance and the conclusions concerning habitat needs as long as there is not a systematic change through time of the microhabitat sites used for calling. The advantage of the microscale is that it measures within mesoscale heterogeneity (e.g., within pond heterogeneity), which cannot be captured easily with the mean prevalence at the mesoscale level. Aggregation reduces the information content but does not make the more rened information obsolete. This is probably also the reason why the microscale has already been identied by others as relevant for biological processes linked to reproduction in amphibians and grasshoppers (Appelt, 1996; Gnther, 1996; Samietz and Berger, 1997; Ingrisch and Khler, 1998). Ponds may not become suitable for amphibian species just because of factors that can be measured only on the pond scale (such as the pond age) but because they have microhabitat features that are avoided, preferred, or essential for amphibians and especially for calling males at different scales. Furthermore, calling males are a key factor in the reproduction and survival of amphibian species. It may be argued that the relative importance of the scales observed by us may be an artefact as they confound with the number of parameters measured at each scale. It is true that for amphibians much less variables were measured at the macroscale than for the remaining two scales. However, the optimal models required only 3e6 variables and only in the case of one species a variable from the macroscale was selected. Moreover, the largest number of variables used in the analysis was at the mesoscale but the microscale provided better prediction accuracy. Also, published uniscale habitat models for amphibian breeding ponds (e.g., Pavignano et al., 1990; Halley et al., 1996), i.e. our mesoscale, required more individual variables to reach acceptable predictive accuracy than our minimum (optimum) models.

5. Conclusions Our study imply trade-offs between efforts in the development of habitat models and their suitability for use in animal conservation, for example in selecting priority sites (Margules and Pressey, 2000; Altmoos and Henle, 2007) or in evaluating and managing the habitats for target species (a common approach in conservation practice). Multi-scale analysis does not make the use of habitat models more complicated for planners and conservationists, as relatively few variables (2 to 6 in our species) will sufce. However, before optimal regional models can be used, a multi-scale approach is needed for their development, and only this may well be more complex than previous uniscale approaches. We have drawn up the following recommendations distinguishing four main aspects (aed): (a) Multi-scaling should always be applied in habitat analyses and modelling Multi-scale analyses should always be carried out for all species in the focus of planning interests. If multi-scaling is not used, important variables for species may be missed out as was shown in our results. (b) Species-specic area partitioning The three nested scales used in our case study are a compromise for modelling; many more scales are conceivable. In any case, judging by our results, it is important to distinguish between at least three different scales: a mesoscale where the majority of analyses are already carried out, below this a microscale, which may be particularly important and is often neglected, and large-area variables at the scale of the local or regional area. This is also a good compromise, which can be used in many practical applications. Three scales were also successfully used in several other studies addressing the effects of scale (e.g., Karl et al., 2000, for birds in Idaho; Luck, 2002, for the Australian rufous treecreeper (C. rufa); Ribas et al., 2005, for insect species and forest fragmentation in Brazil; Caruso et al., 2005, for soil fauna diversity of the Pelagian Islands, Mediterranean). Decisions about the appropriate scales of investigation have to be based on the relevant ecological traits of the species of interest, especially home range size, dispersal ability, and dependence on different habitat types (Morris, 1987; Scott et al., 2002). (c) Special attention to microscales Since the lowest scale proved to be very important in all species investigated, this scale should receive more attention than in the past. (d) Minimum models as suitable trade-offs Minimum models should be derived using an optimal combination of variables from different scales. They represent tradeoffs between the accuracy of information and simplicity and are therefore the ones, which should preferably be drawn up and used in practice (e.g., Carter et al., 2006). These trade-offs are the original idea of habitat modelling in general (see introduction). Multi-scale approaches conrm this concept. Minimum models show those spatial scales that can be disregarded in practice and those which cannot.

Acknowledgement The writing of the manuscript was supported, in part, by the EUproject SCALES (grant #: 226852).

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Appendix. Sampling sizes and measured habitat variables (see Materials and methods)
A: Overview e site sampling. Region: Basic set: Basic set for habitat modelling: Region of approx. 30 30 km south of Leipzig (Germany) 128 Mining sites with different sizes within the region 4 of the 128 Sites

B: Overview e species data set: amphibians. Basic set Sampling objects 199 Ponds in 4 mining sites, 97 of the 199 ponds with at least one amphibian species


and

Calling males (presence) at any pond calling places

Occupied places: specics Unoccupied places

 Habitat variables on 3 scales around the calling places Bufo viridis: 188 calling Pelobates fuscus: 513 calling places at 34 ponds places at 62 ponds Absence of species: 400 places distributed across the 199 ponds

Hyla arborea: 213 calling places at 46 ponds

C: Overview e species data set: grasshoppers. Basic set Sampling objects Unoccupied places (randomly distributed across the sites) 4 Mining sites 1000 Individuals of Sphingonotus caerulans 1000

1000 Individuals of Oedipoda caerulescens 1000

D: Habitat variables and habitat data set: amphibians. Method pH Direct measure with pH-instrument Direct measure with conductivity-instrument Direct measure with N-instrument Presence/absence-survey Field survey and map analysis Visual classication Direct measure 1-Microscale (5 m radius around calling place) 1 Measuring point, ordinal classes (pH 1e2, 2e3, 3e4,.) 1 Measuring point, ordinal classes 1 Measuring point, ordinal classes e e 2 Classes: high e low 5 Classes: 0e0.1 m, 0.1e0.5 m, 0.5e1 m, 1e2 m, >2 m 20% Classes Yeseno Yeseno 20 Intervals 10% Intervals 10% Intervals 10% Intervals Type of soil and classication in 20%-intervals e 2-Mesoscale (pond) Average of all measuring points in the pond, ordinal classes (pH 1e2, 2e3, 3e4,.) Average of all measuring points in the pond, ordinal classes Average of all measuring points in the pond, ordinal classes Presence/absence Classes: <1 year, 1e2 years, 3e5 years, 6e10 years, >10 years 2 Classes: high e low 5 Classes: 0e0.1 m, 0.1e0.5 m, 0.5e1 m, 1e2 m, >2 m 20% Classes Yeseno Yeseno 20 Intervals 10% Intervals 10% Intervals 10% Intervals Type of soil and classication in 20%-intervals 5 Classes: <10 m2, 10e100 m2, 100e1000 m2, 1000e10,000 m2, >10,000 m2 Coverage of habitat types e <100 m, 100 m-intervals, >1000 m 3-Macroscale (landscape) e

Conductivity Nitrogen, total Presence of sh Age Warming capacity Maximum depth

e e e e e e

Fluctuation of water level Turbidity Water ow Slope of bank (inclination) Maximum degree of shading Coverage of submerged vegetation Coverage of emergent vegetation Composition of soil types

Measure of water throughout the year Visual classication Visual classication Inclination classication (maximum) Visual classication (maximum) Visual classication (maximum) Visual classication (maximum) Types of soil (eld survey)

e e e e e e e Type of soil and classication in 20%-intervals e

Pond size

Mapping

Habitat types in surrounding landscapes Number of ponds in areas (300 m, 3 km radius) Distance to next pond

Mapping of habitat types Mapping of ponds Mapping of ponds

Coverage of habitat types e <100 m, 100 m-intervals, >1000 m

Coverage of habitat types, distance to pond 6 Classes: no, 1e5, 6e10, 11e15, 16e20, >20 <100 m, 100 m-intervals, >1000 m

M. Altmoos, K. Henle / Acta Oecologica 36 (2010) 548e560 E: Habitat variables and habitat data set: grasshoppers. Method Total vegetation coverage Vegetation coverage 5 cm above ground Vegetation coverage 20 cm above ground Vegetation coverage 50 cm above ground Percentage of tree shading Percentage of bare ground Soil type Visual classication (coverage maximum) Visual classication (coverage maximum) Visual classication (coverage maximum) Visual classication (coverage maximum) Visual classication (coverage maximum) Visual classication (coverage maximum) Types of soil (survey) 1-Microscale (1 m-radius around individual place) 10% Intervals 10% Intervals 10% Intervals 10% Intervals 10% Intervals e Type of soil and classication in 20%-intervals 20 Intervals Mainly North, East, South, West e e 2-Mesoscale (25 m2 grid) 10% Intervals 10% Intervals 10% Intervals 10% Intervals 10% Intervals 10% Intervals Type of soil and classication in 20%-intervals 20 Intervals Mainly North, East, South, West 3 Classes: no, class 1: 1e5 nets, class 2: 6e10 nets, class 3: >10 nets 5 Classes: class 1: new (<1 year), class 2: 1e2 years, class 3: 3e5 years, class 4: 6e10 years, class 5: >10 years) 3-Macroscale (5000 m2 area) 10% Intervals e e e 10% Intervals 10% Intervals Type of soil and classication in 20%-intervals e e e

559

Slope Exposition (if slope > 20 ) Orb spider abundance class Age

Inclination classication (maximum) Mapping Field survey Field survey and map analysis

5 Classes: class 1: new (<1 year), class 2: 1e2 years, class 3: 3e5 years, class 4: 6e10 years, class 5: >10 years)

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