Effects of historical and present fragmentation on plant species diversity in semi-natural grasslands in Swedish rural landscapes

Habitat loss and fragmentation of natural and semi-natural habitats are considered as major threats to plant species richness. Recently several studies have pinpointed the need to analyse past landscape patterns to understand effects of fragmentation, as the response to landscape change may be slow in many organisms, plants in particular. We compared species richness in continuously grazed and abandoned grasslands in different commonplace rural landscapes in Sweden, and analysed effects of isolation and area in three time-steps (100 and 50 years ago and today). Old cadastral maps and aerial photographs were used to analyse past and present landscape patterns in 25 sites. Two plant diversity measures were investigated; total species richness and species density. During the last 100 years grassland area and connectivity have been reduced by about 90%. Present-day habitat area was positively related to total species richness in both habitats. There was also a relationship to habitat area 50 years ago for continuously grazed grasslands. Only present management was related to species density: continuously grazed grasslands had the highest species density. There were no relationships between grassland connectivity, present or past, and any diversity measure. We conclude that landscape history is not directly important for present-day plant diversity patterns in ordinary landscapes, although past grassland management is a prerequisite for the grassland habitats that can be found there today. It is important that studies are conducted, not only in very diverse landscapes, but also in managed landscapes in order to assess the effects of fragmentation on species.     

Analysis of land-cover transitions based on 17th and 18th century cadastral maps and aerial photographs

This paper explores the possibility of using non-geometric cadastral maps from the 17th and 18th century together with aerial photographs from 1945 and 1981 to analyse land-cover change in south-east Sweden. Habitats rich in plant species in the European rural landscape seem to be correlated with a long continuity of management. Accurate spatial data from historical data sources are fundamental to understand patterns of vegetation and biodiversity in the present-day landscape. However, traditional methods for rectification of non-geometric maps using corresponding points from orthophotos or modern maps are not satisfying, as internal inaccuracies will remain in the maps. This study presents a method to rectify the maps by local warping, thereby eliminating geometrical irregularities. Further, the land-cover changes were calculated and presented as transition matrices. The extent of arable fields and grasslands were analysed in relation to soil characteristics and continuity of management. The results show a dynamic relation between grassland and arable field, albeit the overall proportions remained almost the same between 17th and 18th centuries: 60% grassland to 32% arable field. The most substantial changes in land-cover were prior to 1945. Today there is 18% grasslands left in the study area, while 56% of the land-cover is arable field. Approximately 8% of present-day land-cover is semi-natural grassland 300 years of age or more. Compared to 300 years ago there is only 1% grassland left on peat and 2% on clay. In contrast, grassland covers associated with bare bedrock have been fairly stable in size. All semi-natural grasslands with a long continuity of management were situated on shallow soils, less than 50 cm depth. The major conclusions from this study are that (i) correctly rectified, old maps are very useful to address questions of land-cover changes in historical time, (ii) general trends in land use over 300 years in this hemi-boreal landscape seem to underestimate the full dynamics of land use change, and (iii) only a small proportion of the semi-natural grassland area had a 300 year continuity of management.     

Landscape history of a calcareous (alvar) grassland in Hanila,western Estonia,during the last three hundred years

The landscape history of the largest calcareous seminatural alvar site (ca. 700 ha) in Estonia, is described with the help of a historical map from 1705 and aerial photographs from 1951, and recent vegetation mapping from 1994–1996. The seminatural, species rich alvar grasslands originate and are maintained by grazing of domestic animals. Three hundred years ago the area was mainly open grassland with sparse shrubs and some fields. Forty years ago the vegetation pattern was similar, with some smaller forests and forest clear-cut areas present. Now, since grazing has ceased for ca. 40 years, only 30% of the area remains as open grassland and 70% as forest. Identification of clusters of field layer vegetation using the program TABORD resulted in 8 clusters, which agreed with the empirically determined community types. The field layer within the young pine forest (up to 20 year old pines) is similar to the open alvar grassland. In older forests, the field layer has already changed. There were no phytosociological differences found between ancient grasslands and grasslands on former arable fields or forest clear-cut areas. Decrease in species richness, compared to open grassland, was most drastic in forests of age 20–40 years where the canopy was most closed. Forests have spread more extensively in areas with deeper soil. The continuation of traditional management (grazing and tree cutting) in alvar grasslands is urgently needed in order to keep seminatural alvar grasslands open. The possibility to restore open grasslands remains as long as there is a pool of grassland species available, especially in younger forests.     

Functional rather than structural connectivity explains grassland plant diversity patterns following landscape scale habitat loss

Context

Functional connectivity is vital for plant species dispersal, but little is known about how habitat loss and the presence of green infrastructure interact to affect both functional and structural connectivity, and the impacts of each on species groups.

Objectives

We investigate how changes in the spatial configuration of species-rich grasslands and related green infrastructure such as road verges, hedgerows and forest borders in three European countries have influenced landscape connectivity, and the effects on grassland plant biodiversity.

Methods

We mapped past and present land use for 36 landscapes in Belgium, Germany and Sweden, to estimate connectivity based on simple habitat spatial configuration (structural connectivity) and accounting for effective dispersal and establishment (functional connectivity) around focal grasslands. We used the resulting measures of landscape change to interpret patterns in plant communities.

Results

Increased presence of landscape connecting elements could not compensate for large scale losses of grassland area resulting in substantial declines in structural and functional connectivity. Generalist species were negatively affected by connectivity, and responded most strongly to structural connectivity, while functional connectivity determined the occurrence of grassland specialists in focal grasslands. Restored patches had more generalist species, and a lower density of grassland specialist species than ancient patches.

Conclusions

Protecting both species rich grasslands and dispersal pathways within landscapes is essential for maintaining grassland biodiversity. Our results show that increases in green infrastructure have not been sufficient to offset loss of semi-natural habitat, and that landscape links must be functionally effective in order to contribute to grassland diversity.

     

Modeling scale-dependent landscape pattern,dispersal, and connectivity from the perspective of the organism

Understanding the impacts of habitat fragmentation on dispersal is an important issue in landscape and conservation ecology. Here I examine the effects of fine- to broad-scale patterns in landscape structure on dispersal success of organisms with differing life-history traits. An individual-based model was used to simulate dispersal of amphibian-like species whose movements were driven by land cover and moisture conditions. To systematically control spatial pattern, a landscape model was created by merging simulated land cover maps with synthetic topographic surfaces. Landscapes varied in topographic roughness and spatial contagion in agriculture and urban land cover. Simulations included three different species types that varied in their maximum potential dispersal distances by 1-, 2-, or 4-fold. Two sets of simulations addressed effects of varying aspects of landscape structure on dispersal success. In the first set of simulations, which incorporated variable distances between breeding patches, dispersal success was lowest for all species types when anthropogenic cover was patchily distributed. In the second set, with interpatch distances held constant as landscape composition varied, dispersal success decreased as anthropogenic cover became spatially contagious. Both sets revealed strong main effects of species characteristics, interpatch distances and landscape composition on dispersal success; furthermore, scale-dependent patterns in land cover and moisture gradients had a stronger effect on longer- than shorter-ranging species types. Taken together, these simulations suggest that heuristic conservation strategies could potentially be developed based on important but limited life history information.     

Patterns of habitat occupancy, genetic variation and predicted movement of a flightless bush cricket, Pholidoptera griseoaptera, in an agricultural mosaic landscape

Habitat fragmentation has been generally regarded detrimental to the persistence of many species, especially those with limited dispersal abilities. Yet, when exactly habitat elements become functionally disconnected very much depends on the dispersal ability of a species in combination with the landscape’s composition in which it occurs. Surprisingly, for many small and ground-walking generalists knowledge at what spatial scale and to what extent landscape structure affects dispersal is very scarce. Because it is flightless, the bush cricket Pholidoptera griseoaptera may be regarded susceptible to fragmentation. We applied habitat occupancy surveys, population genetic analyses and movement modelling to investigate the performance of P. griseoaptera in an agricultural mosaic landscape with suitable habitat patches of varying size and isolation. Despite its presumed dispersal limitation we could show that P. griseoaptera occupied the majority of suitable habitats, including small and isolated patches, showed a very low and non-significant genetic differentiation (F _ST = 0.0072) and, in the model, managed to colonize around 73% of all suitable habitat patches within one generation under weak and strong landscape-effect scenarios. We conclude that P. griseoaptera possesses the behavioural attributes (frequent inter-patch dispersal) necessary to persist in this landscape characterized by a patchy distribution of habitat elements. Yet, sound recommendations to landscape planning and conservation require more research to determine whether this represents a general behaviour of the species or a behavioural adaptation to this particular landscape.     

The landscape matrix modifies the effect of habitat fragmentation in grassland butterflies

The landscape matrix is suggested to influence the effect of habitat fragmentation on species richness, but the generality of this prediction has not been tested. Here, we used data from 10 independent studies on butterfly species richness, where the matrix surrounding grassland patches was dominated by either forest or arable land to test if matrix land use influenced the response of species richness to patch area and connectivity. To account for the possibility that some of the observed species use the matrix as their main or complementary habitat, we analysed the effects on total species richness and on the richness of grassland specialist and non-specialist (generalists and specialists on other habitat types) butterflies separately. Specialists and non-specialists were defined separately for each dataset. Total species richness and the richness of grassland specialist butterflies were positively related to patch area and forest cover in the matrix, and negatively to patch isolation. The strength of the species-area relationship was modified by matrix land use and had a slope that decreased with increasing forest cover in the matrix. Potential mechanisms for the weaker effect of grassland fragmentation in forest-dominated landscapes are (1) that the forest matrix is more heterogeneous and contains more resources, (2) that small grassland patches in a matrix dominated by arable land suffer more from negative edge effects or (3) that the arable matrix constitutes a stronger barrier to dispersal between populations. Regardless of the mechanisms, our results show that there are general effects of matrix land use across landscapes and regions, and that landscape management that increases matrix quality can be a complement to habitat restoration and re-creation in fragmented landscapes.     

A multi-scaled analysis of avian response to habitat amount and fragmentation in the Canadian dry mixed-grass prairie

Previous research has suggested that ducks and songbirds may benefit from prairie landscapes that consist primarily of contiguous grasslands. However, the relative importance of landscape-level vs. local characteristics on mechanisms underlying observed patterns is unclear. We measured effects of grassland amount and fragmentation on upland and wetland songbird and duck density and nest success, and on some nest predators, across 16 landscapes in southern Alberta, Canada. We compared these landscape-level effects with local-scale responses, including distance to various edges and vegetation characteristics. We also evaluated several statistical approaches to comparing effects of habitat characteristics at multiple spatial scales. Few species were influenced by grassland amount or fragmentation. In contrast, distance to edge and local vegetation characteristics had significant effects on densities and nest success of many species. Previous studies that reported effects of landscape characteristics may have detected patterns driven by local mechanisms. As a corollary, results were very sensitive to statistical model structure; landscape level effects were much less apparent when local characteristics were included in the models.     

Landscape structure mediates the effects of a stressor on field vole populations

Spatio-temporal landscape heterogeneity has rarely been considered in population-level impact assessments. Here we test whether landscape heterogeneity is important by examining the case of a pesticide applied seasonally to orchards which may affect non-target vole populations, using a validated ecologically realistic and spatially explicit agent-based model. Voles thrive in unmanaged grasslands and untreated orchards but are particularly exposed to applied pesticide treatments during dispersal between optimal habitats. We therefore hypothesised that vole populations do better (1) in landscapes containing more grassland and (2) where areas of grassland are closer to orchards, but (3) do worse if larger areas of orchards are treated with pesticide. To test these hyposeses we made appropriate manipulations to a model landscape occupied by field voles. Pesticide application reduced model population sizes in all three experiments, but populations subsequently wholly or partly recovered. Population depressions were, as predicted, lower in landscapes containing more unmanaged grassland, in landscapes with reduced distance between grassland and orchards, and in landscapes with fewer treated orchards. Population recovery followed a similar pattern except for an unexpected improvement in recovery when the area of treated orchards was increased. Outside the period of pesticide application, orchards increase landscape connectivity and facilitate vole dispersal and so speed population recovery. Overall our results show that accurate prediction of population impact cannot be achieved without taking account of landscape structure. The specifics of landscape structure and habitat connectivity are likely always important in mediating the effects of stressors.     

Determinants of actual functional connectivity for calcareous grassland communities linked by rotational sheep grazing

In fragmented landscapes, plant species persistence depends on functional connectivity in terms of pollen flow to maintain genetic diversity within populations, and seed dispersal to re-colonize habitat patches following local extinction. Connectivity in plants is commonly modeled as a function of the physical distance between patches, without testing alternative dispersal vectors. In addition, pre- and post-dispersal processes such as seed production and establishment are likely to affect patch colonization rates. Here, we test alternative models of potential functional connectivity with different assumptions on source patch effects (patch area and species occupancy) and dispersal (relating to distance among patches, matrix composition, and sheep grazing routes) against empirical patch colonization rates at the community level (actual functional connectivity), accounting for post-dispersal effects in terms of structural elements providing regeneration niches for establishment. Our analyses are based on two surveys in 1989 and in 2009 of 48 habitat specialist plants in 62 previously abandoned calcareous grassland patches in the Southern Franconian Alb in Bavaria, Germany. The best connectivity model S _i , as identified by multi-model inference, combined distance along sheep grazing routes including consistently and intermittently grazed patches with mean species occupancy in 1989 as a proxy for pre-dispersal effects. Community-level patch colonization rates depended to equal degrees on connectivity and post-dispersal process. Our study highlights that actual functional connectivity of calcareous grassland communities cannot be approximated by structural connectivity based on physical distance alone, and modeling of functional connectivity needs to consider pre- and post-dispersal processes.     

Migration rates of grassland plants along corridors in fragmented landscapes assessed with a cellular automation model

This study investigated the efficacy of linear landscape elements in fragmented landscapes as corridors for perennial grassland species with short-range seed dispersal. Corridors are assumed to be essential for the persistence of metapopulations in fragmented landscapes, but it is unclear to what extent linear landscape elements such as ditch banks and road verges can function as corridors for those species. The principal factors that determine the rate of migration through corridors include the width and habitat quality of patches within a corridor (expressed as the population growth rate λ) and the dispersal capacity of plants (expressed as the slope α of the relationship between seed number and log-distance). A cellular automation model was used to simulate the effects of the principal factors on the rate of migration. Simulations with different levels of the principal factors showed highly significant and positive main effects of dispersal capacity, habitat quality and width of corridors on the migration rate. Significant interactions existed between dispersal capacity × width and dispersal capacity × habitat quality (p<0.0001), indicating that the effects of width and habitat quality depended on the dispersal capacity. In narrow corridors most of the dispersed seeds were deposited outside the corridor, which significantly reduced migration rates, especially for species with long-range dispersal (α=−0.4). In wide corridors (up to 20 m), seed losses were much smaller and migration rates approximated those of continuous habitats. The contribution of the few long-range seeds to the rate of migration was significant when habitat quality was high (population growth rates up to 2.5). However, in all simulations migration rates were very low,i.e.<5 m/yr. It is concluded that linear landscape elements are not effective corridors in fragmented landscapes for plants with short-range seed dispersal, because migration rates are low (<5 m/yr), landscape elements vary in the percentage of high quality patches, and refugia and suitable habitat patches are frequently several kilometres apart, making a cohesive infrastructure of corridors for plants elusive. It is argued that the best way to conserve endangered plant species that encounter dispersal barriers is to harvest seeds from nearby source populations and introduce them as suitable habitats.     

Interactive effects of area and connectivity on the diversity of tachinid parasitoids in highly fragmented landscapes

Although many empirical and theoretical studies have elucidated the effects of habitat fragmentation on the third trophic level, little attention has been paid to the impacts of this driver on more generalist groups of non-hymenopteran parasitoids. Here, we used the highly-diverse group of tachinid flies as an alternative model to test the effects of landscape fragmentation on insect parasitoids. Our aims were: (i) to evaluate the relative importance of habitat area and connectivity losses and their potential interaction on tachinid diversity, (ii) to test whether the effects of habitat fragmentation changes seasonally, and (iii) to further assess the effect of habitat diversity on tachinid diversity and whether different parasitoid-host associations modify the species richness response to fragmentation. In 2012 a pan-trap sampling was conducted in 18 semi-natural grasslands embedded in intensive agricultural landscapes along statistically orthogonal gradients of habitat area, connectivity and habitat diversity. We found an interaction between habitat area and connectivity indicating that tachinid abundance and species richness were more negatively affected by habitat loss in landscapes with low rather than with relatively large habitat connectivity. Although tachinid communities exhibited large within-year species turnover, we found that the effects of landscape fragmentation did not change seasonally. We found that habitat diversity and host association did not affect tachinid species diversity. Our results have important implications for biodiversity conservation as any attempts to mitigate the negative effects of habitat loss need to take the general level of habitat connectivity in the landscape into account.     

Multi-scale models of grassland passerine abundance in a fragmented system in Wisconsin

Fragmentation of grasslands has been implicated in grassland bird population declines. Multi-scale models are being increasingly used to assess potential factors that influence grassland bird presence, abundance, and productivity. However, studies rarely assess fragmentation metrics, and seldom evaluate more than two scales or interactions among scales. We evaluated the relative importance of characteristics at multiple scales to patterns in relative abundance of Savannah Sparrow (Passerculus sandwichensis), Grasshopper Sparrow (Ammodramus savannarum), Eastern Meadowlark (Sturnella magna), and Bobolink (Dolichonyx oryzivorus). We surveyed birds in 74 southwestern Wisconsin pastures from 1997 to 1999 and compared models with explanatory variables from multiple scales: within-patch vegetation structure (microhabitat), patch (macrohabitat), and three landscape extents. We also examined interactions between macrohabitat and landscape factors. Core area of pastures was an important predictor of relative abundance, and composition of the landscape was more important than configuration. Relative abundance was frequently higher in pastures with more core area and in landscapes with more grassland and less wooded area. The direction and strength of the effect of core pasture size on relative abundance changed depending on amount of wooded area in the landscape. Relative abundance of grassland birds was associated with landscape variables more frequently at the 1200-m scale than at smaller scales. To develop better predictive models, parameters at multiple scales and their interactive effects should be included, and results should be evaluated in the context of microhabitat variability, landscape composition, and fragmentation in the study area.     

Building patterns and landscape fragmentation in northern Wisconsin,USA

Housing growth is prevalent in rural areas in the United States and landscape fragmentation is one of its many effects. Since the 1930s, rural sprawl has been increasing in areas rich in recreational amenities. The question is how housing growth has affected landscape fragmentation. We thus tested three hypotheses relating land cover and land ownership to density and spatial pattern of buildings, and examined whether building density or spatial pattern of buildings was a better predictor for landscape fragmentation. Housing locations were mapped from 117 1:24,000-scale USGS topographic maps across northern Wisconsin. Patch-level landscape metrics were calculated on the terrestrial area remaining after applying 50, 100 and 250 m disturbance zones around each building. Our results showed that building density and the spatial pattern of buildings were affected mostly by lake area, public land ownership, and the abundance of coniferous forest, agricultural land, and grassland. A full 40% of the houses were within 100 m of lakeshores. The clustering of buildings within 100 m of lakeshores limited fragmentation farther away. In contrast, agricultural and grassland areas were correlated with higher building density, higher fragmentation, and more dispersed building pattern possible legacies of agricultural settlement patterns. Understanding which factors influence building density and fragmentation is useful for landscape level planning and ecosystem management in northern Wisconsin and areas that share similar social and environmental constraints.     

Historic landscape change and habitat loss: the case of black grouse in Lower Saxony,Germany

The declines of many specialist bird species in the agricultural landscapes of Central Europe have resulted in small and isolated populations. In the case of the black grouse, a ground-nesting bird species with large spatial requirements, empiric evidence about underlying landscape changes is scarce. In this study, we examined land cover and land cover changes in a farmland-forest mosaic in eastern Lower Saxony, Germany and how they affect occurrence and persistence of black grouse. Spatial information came from historic topographic maps from 1958 to 1975. The results show profound conversions of habitat to forest and farmland but also an increase in settlement area. Habitat conversions and suburbanization were negative correlates of black grouse persistence. Habitat models from before and after a decline period differed in some of the predictors and suggest black grouse habitat to be more diverse before the land cover changes. Our study confirms that land use factors at a landscape scale extent contribute to explain black grouse occurrence and thus can complement important small scale factors like the quality and size of individual habitat patches. Results also show that landscape factors affect black grouse distribution predominantly from an area much greater than an individual black grouse home range. Our models may be further evaluated on present-day landscapes and might be used to evaluate large-scale habitat availability for black grouse.     

Landscape and habitat filters jointly drive richness and abundance of specialist plants in terrestrial habitat islands

Context

Landscape and habitat filters are major drivers of biodiversity of small habitat islands by influencing dispersal and extinction events in plant metapopulations.

Objectives

We assessed the effects of landscape and habitat filters on the species richness, abundance and trait composition of grassland specialist and generalist plants in small habitat islands. We studied traits related to functional spatial connectivity (dispersal ability by wind and animals) and temporal connectivity (clonality and seed bank persistence) using model selection.

Methods

We sampled herbaceous plants, landscape (local and regional isolation) and habitat filters (inclination, woody encroachment and disturbance) in 82 grassland islands in Hungary.

Results

Isolation decreased the abundance of good disperser specialist plants due to the lack of directional vectors transferring seeds between suitable habitat patches. Clonality was an effective strategy, but persistent seed bank did not support the survival of specialist plants in isolated habitats. Generalist plants were unaffected by landscape filters due to their wide habitat breadth and high propagule availability. Clonal specialist plants could cope with increasing woody encroachment due to their high resistance against environmental changes; however, they could not cope with intensive disturbance. Steep slopes providing environmental heterogeneity had an overall positive effect on species richness.

Conclusions

Specialist plants were influenced by the interplay of landscape filters influencing their abundance and habitat filters affecting species richness. Landscape filtering by isolation influenced the abundance of specialist plants by regulating seed dispersal. Habitat filters sorted species that could establish and persist at a site by influencing microsite availability and quality.

     

How do local habitat management and landscape structure at different spatial scales affect fritillary butterfly distribution on fragmented wetlands?

Habitat fragmentation, patch quality and landscape structure are important predictors for species richness. However, conservation strategies targeting single species mainly focus on habitat patches and neglect possible effects of the surrounding landscape. This project assesses the impact of management, habitat fragmentation and landscape structure at different spatial scales on the distribution of three endangered butterfly species, Boloria selene, Boloria titania and Brenthis ino. We selected 36 study sites in the Swiss Alps differing in (1) the proportion of suitable habitat (i.e., wetlands); (2) the proportion of potential dispersal barriers (forest) in the surrounding landscape; (3) altitude; (4) habitat area and (5) management (mowing versus grazing). Three surveys per study site were conducted during the adult flight period to estimate occurrence and density of each species. For the best disperser B. selene the probability of occurrence was positively related to increasing proportion of wetland on a large spatial scale (radius: 4,000 m), for the medium disperser B. ino on an intermediate spatial scale (2,000 m) and for the poorest disperser B. titania on a small spatial scale (1,000 m). Nearby forest did not negatively affect butterfly species distribution but instead enhanced the probability of occurrence and the population density of B. titania. The fen-specialist B. selene had a higher probability of occurrence and higher population densities on grazed compared to mown fens. The altitude of the habitat patches affected the occurrence of the three species and increasing habitat area enhanced the probability of occurrence of B. selene and B. ino. We conclude that, the surrounding landscape is of relevance for species distribution, but management and habitat fragmentation are often more important. We suggest that butterfly conservation should not focus only on a patch scale, but also on a landscape scale, taking into account species-specific dispersal abilities.     

Interactions between plant life span, seed dispersal capacity and fecundity determine metapopulation viability in a dynamic landscape

Classical metapopulation models do not account for temporal changes in the suitability of habitat patches. In reality, however, the carrying capacity of most habitat types is not constant in time due to natural succession processes. In this study, we modeled plant metapopulation persistence in a successional landscape with disappearing and emerging habitat patches, based on a realistic dune slack landscape at the Belgian–French coast. We focused on the effects of the variation of different plant traits on metapopulation persistence in this changing landscape. Therefore, we used a stage based stochastic metapopulation model implemented in RAMAS/Metapop, simulating a large variation in plant traits but keeping landscape characteristics such as patch turnover rate and patch lifespan constant. The results confirm the conclusions of earlier modeling work that seed dispersal distance and seed emigration rate both have an important effect on metapopulation persistence. We also found that high population growth rate or high recruitment considerably decreased the extinction risk of the metapopulation. Additionally, a long plant life span had a strong positive effect on metapopulation persistence, irrespective of the plant's dispersal capacity and population growth rate. Plant species that invest in life span require less investment in offspring and dispersal capacity to avoid extinction, even in dynamic landscapes with deterministic changes in habitat quality. Moreover, metapopulations of long-lived plant species were found to be much less sensitive to high levels of environmental stochasticity than short-lived species.     

Testing coexistence of extinction debt and colonization credit in fragmented calcareous grasslands with complex historical dynamics

Calcareous grasslands are among the most species-rich ecosystems in temperate countries. However, these ecosystems have suffered from fragmentation and destruction during the last century. We studied the response of calcareous grassland plant diversity to landscape changes in Belgium. Results indicated that high area loss (since 1965) old habitat patches exhibited an extinction debt inverse to low area loss old habitat patches, little depending on the area loss threshold (60%, 70%, 80% or 90%) considered for the distinction between the high and low area loss patches. However, human activities also created new habitat patches in the landscape and therefore provided opportunities for calcareous grassland plant species to colonize new habitats. This also provided opportunities to study species colonization abilities in the context of habitat restoration. We analyzed species richness in new patches compared to old patches in order to detect colonization credit. We detected the presence of a colonization credit in new patches when using high loss old patches (area loss >80%, exhibiting an extinction debt) or all old patches as a reference. However, when the reference was low loss old patches alone (area loss <80%, less likely to exhibit an extinction debt), no colonization credit was detected. In addition, species composition was similar between new patches and old patches. These results are encouraging for restoration programs. However, the results indicated that the presence of an extinction debt in reference habitats could lead to inaccurate conclusions in restoration monitoring. Therefore, extinction debt should be considered when choosing reference habitats to evaluate restoration success.