‘冬枣’天然二、四混倍体的发现与评价

以山东省沾化县栽培的普通‘冬枣’及其3个大果型变异类型‘沾冬2号’、变异1号、变异2号为试材,对其染色体倍性、DNA指纹、果实和叶片形态及果实主要营养指标进行了比较分析。经流式细胞仪测定,3个大果型变异类型均为二、四混倍体（2x + 4x）,其对照（母株）均为纯二倍体。经74对SSR引物检测,未发现3个大果型混倍体变异与其二倍体对照的DNA指纹存在差异,表明三者均确系来源于普通的二倍体‘冬枣’,其四倍体部分均属于同源加倍。与普通‘冬枣’相比,3个变异类型均表现出叶片变宽,叶形指数变小,果实变大,果形指数变小,果实维生素C含量增加,但果实可溶性糖和可滴定酸含量变化无规律。与已通过良种审定的‘沾冬2号’相比,新发现的变异1号和变异2号的单果质量更大,分别增大22.95%和15.90%,分别达到27.05 g和25.50 g。     

Spatial configuration matters: a test of the habitat amount hypothesis for plants in calcareous grasslands

Context

A recent hypothesis, the habitat amount hypothesis, predicts that the total amount of habitat in the landscape can replace habitat patch size and isolation in studies of species richness in fragmented landscapes.

Objectives

To test the habitat amount hypothesis by first evaluating at which spatial scale the relationship between species richness in equal-sized sample quadrats and habitat amount was the strongest, and then test the importance of spatial configuration of habitat—measured as local patch size and isolation—when habitat amount was taken into account.

Methods

A quasi-experimental setup with 20 habitat patches of dry calcareous grasslands varying in patch size, patch isolation and habitat amount at the landscape scale was established in the inner Oslo fjord, Southern Norway. We recorded species richness of habitat specialists of vascular plants in equal-sized sample quadrats and analysed the relationship between species richness, habitat amount in the landscape and patch size and isolation.

Results

Although the total amount of habitat in a 3 km-radius around the local patch was positively related to species richness in the sample quadrats, local patch size had an additional positive effect, and the effect of patch size was higher when the amount of habitat within the 3 km-radius was high than when it was low.

Conclusions

In our study system of specialist vascular plants in dry calcareous grasslands, we do not find support for the habitat amount hypothesis.

     

How does the landscape context of occurrence data influence models of invasion risk? A comparison of independent datasets in Massachusetts,USA

The spatial distribution of non-native, invasive plants on the landscape is strongly influenced by human action. People introduce non-native species to new landscapes and regions (propagule pressure) as well as increase ecosystem invasibility through disturbance of native ecosystems. However, the relative importance of different landscape drivers of invasion may vary with landscape context (i.e., the types and amounts of surrounding land cover and land use). If so, data collected in one context may not be appropriate for predicting invasion risk across a broader landscape. To test whether independent occurrence datasets suggest similar landscape drivers of invasion, we compared landscape models based on data compiled by the Invasive Plant Atlas of New England (IPANE), which are contributed opportunistically by trained citizen scientists, to models based on Forest Stewardship plans (FSPs), which are located in privately owned and relatively undisturbed forests. We evaluated 16 landscape variables related to propagule pressure and/or disturbance for significant predictors of invasive plant presence based on presence/absence and count regression models. Presence and richness of invasive plants within FSPs was most influenced by proportion of open land and proximity to residential areas, which are both sources of propagules in forest interiors. In contrast, IPANE invasive plant presence and richness for the same area was influenced by distance to roads and streams. These results suggest that landscape drivers of invasion vary considerably depending on landscape context, and the choice of occurrence dataset will strongly influence model results.     

Scale dependence of tree abundance and richness in a tropical rain forest,Malaysia

Abundance and richness are the two fundamental components of speciesdiversity. They represent two distinct types of variables of which the formerisadditive when aggregated across scales while the latter is nonadditive. Thisstudy investigated the changes in the spatial patterns of abundance andrichnessof tree species across multiple scales in a tropical rain forest of Malaysiaandtheir variations in different regions of the study area. The results showedthatfrom fine to coarse scales abundance had a gradual and systematic change inpattern, whereas the change in richness was much less predictable and ahotspot in richness at one scale may become acoldspot at another. The study also demonstrated that differentmeasures of diversity variation (e.g., variance and coefficient of variation)can result in different or even contradictory results which further complicatedthe interpretation of diversity patterns. Because of scale effect the commonlyused measure of species diversity in terms of unit area (e.g.,species/m²) is misleading and of little use in comparing speciesdiversitybetween different ecosystems. Extra care must be taken if management andconservation of species diversity have to be based on information gathered at asingle scale.This revised version was published online in May 2005 with corrections to the Cover Date.     

Epiphyte communities in Mediterranean fragmented forests: importance of the fragment size and the surrounding matrix

Context

Mediterranean forests have been fragmented intensively over time, thereby yielding small and isolated forest remnants. They host a rich variety of epiphytes, which may be affected by landscape structure. Previous studies have analyzed the influence of habitat quality on these epiphytic communities, but there is little knowledge of the effects of other fragment features.

Objectives

We evaluated the impacts of forest loss and fragmentation on epiphytic communities (lichens and bryophytes) at plot and fragment scales after controlling the variation in forest structure and management.

Methods

We considered 40 fragments of dense oak forests in a human-modified landscape. We quantified their spatial attributes (size and shape), the quality of the surrounding matrix and the forest stand structure. We modeled community traits, and the presence and abundance of species at fragment and plot scales.

Results

Fragment size, shape, and the quality of the surrounding matrix were key factors that affected epiphytic richness and diversity. Larger and more regularly shaped fragments hosted the richest and most diverse communities, possibly offering a larger core area and thus favoring the entry of typical forest species. A high-contrast matrix was only favorable in small fragments, probably allowing the arrival of propagules. The species-level response was highly variable.

Conclusions

Landscape structure provides powerful explanations of the richness and diversity losses among epiphytes. Forest management should ensure the retention of the largest possible continuous forests. The management strategy of the matrix will depend on the conservation goal, since we observed different effects related with quality and fragment size.

     

Species richness and composition differ in response to landscape and biogeography

Context

Understanding how landscape patterns affect species diversity is of great importance in the fields of biogeography, landscape ecology and conservation planning, but despite the rapid advance in biodiversity analysis, investigations of spatial effects on biodiversity are still largely focused on species richness.

Objectives

We wanted to know if and how species richness and species composition are differentially driven by the spatial measures dominating studies in landscape ecology and biogeography. As both measures require the same limited presence/absence information, it is important to choose an appropriate diversity measure, as differing results could have important consequences for interpreting ecological processes.

Methods

We recorded plant occurrences on 112 islands in the Baltic archipelago. Species richness and composition were calculated for each island, and the explanatory power of island area and habitat heterogeneity, distance to mainland and structural connectivity at three different landscape sizes were examined.

Results

A total of 354 different plant species were recorded. The influence of landscape variables differed depending on which diversity measure was used. Island area and structural connectivity determined plant species richness, while species composition revealed a more complex pattern, being influenced by island area, habitat heterogeneity and structural connectivity.

Conclusions

Although both measures require the same basic input data, species composition can reveal more about the ecological processes affecting plant communities in fragmented landscapes than species richness alone. Therefore, we recommend that species community composition should be used as an additional standard measure of diversity for biogeography, landscape ecology and conservation planning.

     

Discrimination of native and exotic forest patterns through shape irregularity indices: An analysis in the landscapes of Galicia,Spain

Landscapes resulting from human activity may be expected to present simpler shapes than more natural landscapes. In the case of forest landscapes, the boundaries of native forest patches may be more irregular than those of exotic forest plantations. There is however a lack of quantitative results to this respect, and it is not clear which shape indices are more adequate for such discrimination. In this study, we analysed the shape of a large number of forest classes in the region of Galicia (Spain) using the Spanish Forest Map at a scale 1:50000 as the spatial information source. We considered a set of fifteen shape irregularity indices including those that have been commonly used in landscape ecology studies. We found systematic differences in the shape of the analysed forest classes, with native forests presenting both more complex and elongated boundaries than exotic forests. We suggest that these differences are due to the combined effects of human action and other topographical and hydrological factors. The only index that perfectly discriminated both types of forest was the mean circumscribing circle index. Other six indices provided also a significantly good discrimination: density of shape characteristic points, area-weighted mean perimeter-area ratio, area-weighted mean contiguity index, mean shape index, perimeter-area fractal dimension and mean largest axis index. Comparisons of pure and mixed forests with the same dominant species indicated that an increase in tree species richness is in general associated with more irregular boundaries in the forest. Discarding indices on the basis of a high statistical correlation may not be an adequate procedure to retain the best-performing indices. Finally, we discussed several limitations of some frequently used indices that may be relevant to prevent an improper characterization of landscape shape.     

Does the response of bird assemblages to fire mosaic properties vary among spatial scales and foraging guilds?

Context

An increasing number of studies have investigated the impact of environmental heterogeneity on faunal assemblages when measured at multiple spatial scales. Few studies, however, have considered how the effects of heterogeneity on fauna vary with the spatial scale at which the response variable is characterised.

Objectives

We investigated the relationship between landscape properties in a region characterised by diverse fire mosaics, and the structure and composition of avian assemblages measured at both the site- (1 ha) and landscape-scale (100 ha).

Methods

We surveyed birds and calculated spatial landscape properties in sub-tropical woodlands of central Queensland, Australia.

Results

Environmental heterogeneity, as measured by topographic complexity, was consistently important for bird species richness and composition. However, the explanatory power of topographic complexity varied depending on the spatial scale and the component of diversity under investigation. We found different correlates of richness within particular foraging guilds depending on the scale at which richness was measured. Extent of long-unburnt habitat (>10 years since fire) was the most important variable for the landscape-scale richness of frugivores, insectivores and canopy feeders, whereas environmental heterogeneity in the surrounding landscape was more important for site-scale richness of these foraging guilds.

Conclusions

The response of species richness to landscape characteristics varies among scales, and among components of diversity. Thus, depending on the scale at which a biodiversity conservation goal is conceptualised—maximising richness at a site, or across a landscape—different landscape management approaches may be preferred.

     

Ecological relationships between landscape change and plant guilds in depressional wetlands

Plant guilds used to measure the relationships between wetlandplant community characteristics and landscape change around 31 depressionalwetlands in central Ohio, USA. Characteristics of certain plant guilds withineach wetland site are correlated with changes in: (a) area of urban land cover,forest, grassland, agriculture, and open-water in the local vicinity ofthe wetland; (b) inter-wetland distance; and (c) wetland size (area).Taxa richness is negatively correlated with inter-wetland distance forall plant guilds, except submersed herbaceous plants. Taxa richness of thesubmersed herbaceous plant guild (usually less than 20% of the totalnumber of plant species at a wetland) is positively correlated with the area ofopen-water in the local landscape and with the areaofthe wetland site itself. Significant positive correlationsalso exist between the area of open-water in the vicinity of the wetlandand the proportion of submersed herbaceous plant taxa at the site, the numberofnative submersed herbaceous plant species, the submersed herbaceous plantperennial-to-annual ratio, and the number ofavian-dispersed submersed herbaceous plant species at a site. Theresults suggest that (a) the dominance of submersed herbaceousplantspecies at a site is related to dispersal constraints between wetlands, and (b)the relatively slower physiological response of woody plants to local landscapechange may result in their contribution to greater ecologicalinertia in the plant community as a whole. For these reasons,relationships between the plant community and land cover change may not alwaysbe observed unless analyzed at the level of plant-guild.This revised version was published online in May 2005 with corrections to the Cover Date.     

Contemporary spatial and environmental factors determine vascular plant species richness on highly fragmented meadows in Central Finland

Context

Habitat loss is a major threat to biodiversity. It can create temporal lags in decline of species in relation to destruction of habitat coverage. Plant species specialized in semi-natural grasslands, especially meadows, often express such extinction debt.

Objectives

We studied habitat loss and fragmentation of meadows and examined whether the changes in meadow coverage had caused an extinction debt on vascular plants. We also studied whether historical or present landscape patterns or contemporary environmental factors were more important determinants of species occurrence.

Methods

We surveyed the plant species assemblages of 12 grazed and 12 mown meadows in Central Finland and detected the meadow coverages from their surroundings on two spatial scales and on three time steps. We modelled the effects of functional connectivity, habitat amount, and isolation on species richness and community composition.

Results

We observed drastic and dynamic meadow loss in landscapes surrounding our study sites during the last 150 years. However, we did not find explicit evidence for an extinction debt in meadow plants. The observed species richness correlated with contemporary factors, whereas both contemporary factors and habitat availability during the 1960s affected community composition.

Conclusions

Effective conservation management of meadow biodiversity builds on accurate understanding of the relative importance of past and present factors on species assemblages. Both mown and grazed meadows with high species richness need to be managed in the future. The management effort should preferably be targeted to sites located near to each other.

     

Determinants of plant species richness and patterns of nestedness in fragmented landscapes: evidence from land-bridge islands

Land-bridge islands formed by dam construction are considered to be “experimental” systems for studying the effects of habitat loss and fragmentation, offering many distinct advantages over terrestrial fragments. The Thousand Island Lake in Southeast China is one such land-bridge system with more than 1000 islands. Based on a field survey of vascular plant richness on 154 land-bridge islands during 2007–2008, we examined the effects of island and landscape attributes on plant species richness and patterns of species nestedness. We also examined the different responses of plant functional groups (classified according to growth form and shade tolerance) to fragmentation. We found that island area explained the greatest amount of variation in plant species richness. Island area and shape index positively affected species diversity and the degree of nestedness exhibited by plant communities while the perimeter to area ratio of the islands had a negative effect. Shade-tolerant plants were the most sensitive species group to habitat fragmentation. Isolation negatively affected the degree of nestedness in herb and shade-intolerant plants including species with various dispersal abilities in the fragmented landscape. Based on these results, we concluded that the effects of habitat loss and fragmentation on overall species richness depended mostly on the degree of habitat loss, but patterns of nestedness were generated from different ecological mechanisms due to species-specific responses to different characteristics of habitat patches.     

Quantifying habitat specificity to assess the contribution of a patch to species richness at a landscape scale

Assessing and predicting the species richness of a complex landscape remains a problem because there is no simple scaling function of species richness in a heterogeneous environment. Furthermore, the potential value of an area for biodiversity conservation may depend on which, rather than how many, species the area contains. This paper shows how we can objectively evaluate the contribution of an area, e.g., a habitat patch, to larger-scale plant species richness, e.g., a landscape composed of patches of several habitat types, and how we can test hypotheses that attempt to explain this contribution. We quantified the concept of habitat specificity to assess the proportion of each observed plant population that is concentrated within a given spatial element. A case study of a biodiversity-monitoring program in the Swiss Canton of Aargau showed that the relative contribution of the three main types of land use to the overall species richness differed strongly between higher taxa (vascular plants and molluscs). However, the type of data, i.e., presence-absence or abundance, was not important. Resampling of the plant data suggested that stratification provided an unbiased estimate of relative specificity, whereas unstratified sampling caused bias even for large samples. In a second case study of vascular plants in an agricultural landscape in central Switzerland, we tested whether the type, size or shape of a landscape element can predict its contribution to the species richness of the landscape. Habitat types that were less frequently disturbed contributed more per m² to landscape species richness than more frequently disturbed ones. Contrary to expectation, patch size was negatively correlated to specificity per m² for arable fields, whereas patch shape appeared to be unrelated to the specificity per m² both for arable fields and for meadows. The specificity approach provides a solution to the problem of scaling species richness and is ideally suited for testing hypotheses on the effect of landscape structure on landscape species richness. Specificity scores can easily be combined with measures of other aspects of rarity to assess the contribution of a spatial element to conservation goals formulated at regional, national or global level.     

The influence of matrix quality on species richness in remnant forest

Context

Habitat destruction is the leading threat to terrestrial biodiversity, isolating remnant habitat in a matrix of modified vegetation.

Objectives

Our goal was to determine how species richness in several broad taxonomic groups from remnant forest was influenced by matrix quality, which we characterized by comparing plant biomass in forest and the surrounding matrix.

Methods

We coupled data on species-area relationships (SARs) in forest remnants from 45 previously published studies with an index of matrix quality calculated using new estimates of plant biomass derived from satellite imagery.

Results

The effect size of SARs was greatest in landscapes with low matrix quality and little forest cover. SARs were generally stronger for volant than for non-volant species. For the terrestrial taxa included in our analysis, matrix quality decreased as the proportion of water, ice, or urbanization in a landscape increased.

Conclusions

We clearly demonstrate that matrix quality plays a major role in determining patterns of species richness in remnant forest. A key implication of our work is that activities that increase matrix quality, such as active and passive habitat restoration, may be important conservation measure for maintaining and restoring biodiversity in modified landscapes.

     

A landscape-level assessment of understory diversity in upland forests of North-Central Wisconsin,USA

The study measured landscape level diversity of the understory plants of mature, upland forests in north-central Wisconsin USA. Habitat types were used to segregate the landscape along a moisture-nutrient gradient. Forty sites that had closed canopies, had been undisturbed for at least 20 years, and were at least 8 ha in size were used. The percent cover of groundlayer species was ocularly estimated in 12–18 randomly located, one meter square plots in June and August, 1995. Shrub cover was estimated by the line intercept method. Alpha, beta and gamma diversity were determined for early and late summer periods separately. Gamma diversity was quantified using a new method, affinity analysis, which generates a list of modal and outlier sites and calculates mosaic diversity, a measure of landscape complexity. Generally, communities in the middle of the moisture-nutrient gradient were modal, whereas those at the mesic end of the gradient were outlier. Mosaic diversity values were very similar for early summer and late summer (2.88±0.04, 2.95±0.03, respectively), but was much higher for both periods combined (3.95±0.07). Whittaker's Index (beta diversity) revealed varying rates of species turnover along presumed moisture and nutrient gradients, whereas species densities and richness were relatively constant among habitat types. A one-way analysis of variance of Shannon-Weaver values found no significant differences among habitat types (p0.05). Regional diversity mainly resulted from high beta values which appears to be primarily a function of the moisture gradient. The other factors influencing compositional differences among sites are variation in site history, especially disturbance, with niche partitioning and differences in seed dispersal capacity having a minor influence. The affinity analysis method indicated that sampling once per season is inadequate, and that many types of sites are modal. This method for estimating gamma (landscape) diversity shows considerable promise, but information on the processes that produce outlier sites is needed to fully understand and use the results of this method.     

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.

     

Characterizing the complexity of landscape boundaries by remote sensing

This paper presents a method for characterizing the complexity of landscape boundaries by remote sensing. This characterization is supported by a new boundary typology, that takes into account points where three or more landcovers converge (i.e., convergency points or coverts). Landscape boundary richness and diversity indices were proposed and calculated over 19 landscapes in South-East Brazil. Results showed that landscape boundaries, especially convergency points, provided an enrichment in landscape pattern analysis. Landcover boundary diversities were significantly related to landcover shape: elongated riparian units had the highest values for boundary diversity and coverts proportion indices. On the other hand, landscape analysis showed that indices of shape, richness, diversity and coverts proportion provided an additional evaluation of landcover spatial distribution within the landscape.     

A mathematical comment on the formulae for the aggregation index and the shape index

Ina recent paper [Landscape Ecol. 15: 591–601 (2000)] He et al. describedanaggregation index AI _i to measure pixelaggregation within a single class i. We show that thecommonly used shape index SI _i is related to theproposed aggregation metric as SI _i =(A _i) +AI _i(1 –(A _i)), with(A _i) dependent on class areaA _i only. Moreover, it is shown that thenormalized shape index, SI _i ^*,equals (1 – AI _i). We conclude thatAI _i does not provide any information notprovided by SI _i, orSI _i ^*.This revised version was published online in May 2005 with corrections to the Cover Date.     

Towards an energy–landscape integrated analysis? Exploring the links between socio-metabolic disturbance and landscape ecology performance (Mallorca,Spain, 1956–2011)

Context

The role of agricultural landscapes in biodiversity conservation is an emerging topic in a world experiencing a worrying decrease of species richness. Farm systems may either decrease or increase biological diversity, depending on land-use intensities and management.

Objectives

We present an intermediate disturbance-complexity model (IDC) of cultural landscapes aimed at assessing how different levels of anthropogenic disturbance on ecosystems affect the capacity to host biodiversity depending on the land matrix heterogeneity. It is applied to the Mallorca Island, amidst the Mediterranean biodiversity hotspot.

Methods

The model uses the disturbance exerted when farmers alter the Net Primary Production through land-use change as well as when they remove a share of it (HANPP), together with Shannon–Wiener index (H′) of land-cover diversity. The model is tested with a twofold-scalar experimental design (1:50,000 and 1:5000) of a set of landscape units along three time points (1956, 1989, 2011). Species richness of breeding and wintering birds, taken as a biodiversity proxy, is used in an exploratory factor analysis.

Results

The results clearly show that when intermediate levels of HANPP are performed within intermediate levels of complexity (H′) in landscape patterns, like agro-forest mosaics, great bird species richness and high socio-ecological resilience can be maintained. Yet, these complex-heterogeneous landscapes are currently vanishing due to industrial farm intensification, rural abandonment and urban sprawl.

Conclusions

The results make apparent the usefulness of transferring the concept of intermediate disturbance-complexity interplay to cultural landscapes. Our spatial-explicit IDC model can be used as a tool for strategic environmental assessment of land-use planning.

     

Diversity–disturbance relationship in forest landscapes

Context

Despite decades of research, there is an intense debate about the consistency of the hump-shaped pattern describing the relationship between diversity and disturbance as predicted by the intermediate disturbance hypothesis (IDH). Previous meta-analyses have not explicitly considered interactive effects of disturbance frequency and intensity of disturbance on plant species diversity in terrestrial landscapes.

Objective

We conducted meta-analyses to test the applicability of IDH by simultaneously examining the relationship between species richness, disturbance frequency (quantified as time since last disturbance as originally proposed) and intensity of disturbance in forest landscapes.

Methods

The effects of disturbance frequency, intensity, and their interaction on species richness was evaluated using a mixed-effects model.

Results

We found that species richness peaks at intermediate frequency after both high and intermediate disturbance intensities, but the richness-frequency relationship differed between intensity classes.

Conclusions

Our study highlights the need to measure multiple disturbance components that could help reconcile conflicting empirical results on the effect of disturbance on plant species diversity.

     

Biodiversity functions of urban cemeteries: Evidence from one of the largest Jewish cemeteries in Europe

As the world becomes more urbanized, urban cemeteries may become increasingly valuable for biodiversity conservation as cemeteries are ubiquitous elements of the green infrastructure in cities worldwide. By implementing a multi-taxon approach at different spatial extents, we analyzed habitat functions of a large urban cemetery in Berlin (Weiÿensee Jewish Cemetery) and explored related environmental variables. This cemetery is an outstanding cultural heritage site but it also stands for old urban cemeteries that have progressed to urban woodland, an ecosystem type that exists in many regional and religious contexts. The cemetery provided a habitat for 604 species; species of conservation concern comprised 1.6⿿100% of total species among different groups of taxa (in decreasing order: bats, birds, lichens, bryophytes, carabids, vascular plants, spiders). Species richness and species composition at the plot level were significantly related to differences in management intensity and resulting vegetation structures but differed among taxonomic groups. In vascular plants, carabids and spiders, the species composition varied significantly with habitat age, and there was a set of characteristic species for different age classes in each species group. Our results thus support the use of differentiated management approaches to maintain habitat heterogeneity by allowing wilderness development in some parts of a cemetery while keeping others more open. Since these aims can be combined with efforts to preserve outstanding grave architectures and allow access to visitors, our study indicates ways of reconciling conflicting aims of heritage preservation and biodiversity conservation, a promising perspective for biodiversity conservation in culturally shaped urban landscapes. We conclude that cemeteries provide important cultural ecosystem services within the urban green infrastructure.