What role can cemeteries play in conserving plant biodiversity in different urbanisation contexts?

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Jeanne Vallet This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7448159/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 09 Apr, 2026 Read the published version in Urban Ecosystems → Version 1 posted 9 You are reading this latest preprint version Abstract The aim of this work is to assess the importance and the originality of plant vascular biodiversity of cemeteries at a regional scale, compare it with other urbanised habitats and assessing their conservation issues according to an urban-rural gradient. The study was carried out in the Île-de-France region (France) with a dataset containing 66,973 records of vascular plants collected in 817 cemeteries between 2000 and 2021. We analysed the preference of species for cemeteries or other anthropogenic habitats using indicator value analysis. To describe species-specific responses to the urban-rural gradient, we employed a niche model analysis coupled with a randomization procedure. We then compared naturalness and habitat preferences between the different species lists obtained by these two methods. Among the 731 spontaneous species were inventoried, 84% of which are native. Of these, 69 can be considered as species of conservation interest; species of anthropogenic communities were well represented but the majority were linked to semi-natural habitats: heathlands, dry grasslands on acidic or calcareous soils and meadows in particular. Species linked to cemeteries are associated with semi-natural habitats, while the species linked to other urban habitats are forest or wetland species. Finally, species specifically linked to cemeteries in an urban context are mainly non-native species. Species of semi-natural herbaceous habitats are much more represented among the species linked to rural cemeteries. When they exist in cemeteries, the conservation of remnant semi-natural habitats should be a priority. remnant habitats nature conservation species of conservation interest non-native species graveyards urban planning Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Introduction Given the extent of land use transformations on a planetary scale due to the development of urbanization and agriculture, it is now essential to also consider the conservation of biodiversity in these anthropogenic spaces (Planchuelo et al. 2019). Different approaches can be envisaged to take better account of biodiversity in these anthropogenic contexts: Planchuelo et al (2019) stress the importance of all types of ecosystem for the conservation of plant species of conservation interest, whether they have been profoundly altered by human activities (novel ecosystems) or are made up of fragments of relict habitats (remnant ecosystems) or a hybrid between the two. Cemeteries are found in almost every town and village; they can vary greatly in terms of size, age, layout, funeral practices and the degree of urbanisation in the surrounding area. Generally speaking, cemeteries fall into the category of new ecosystems (Planchuelo et al. 2019), but some cemeteries have the particularity of harbouring remant habitats that may be of great interest for the conservation of plant species (Löki et al. 2020; Molnár et al. 2017). Cemeteries have also been identified as a potential source of non-native species, some of which may be invasive (Walusiak and Krztoń 2021). Many studies on cemetery flora have focused on a few cemeteries that are often singular in terms of their characteristics (Buldrini et al. 2023; Yılmaz et al. 2018; Yılmaz et al. 2018) and few have taken a more global approach on the scale of an entire territory (Buch et al. 2025). Moreover, as funerary practices can vary widely between countries, religions etc., the results cannot necessarily be more broadly generalised. Few studies have been conducted on the biodiversity of cemeteries in Western Europe (Itescu and Jeschke 2024). Finally, the 2020s mark a pivotal period in terms of management practices for cemeteries in France (European Environment Agency 2023) because since the 1 st of July 2022, pesticides are prohibited in these areas 1 ; a number of municipalities have anticipated the law (Flandin 2019). It is therefore particularly important to identify the state of present biodiversity in cemeteries in order to understand the future effects of this change in management. The aim of this study is to examine the vascular plant biodiversity of cemeteries in Île-de-France, a region surrounding Paris where human pressure is particularly strong, with agricultural areas occupying around 50% of the regional surface area and urban areas 23% (Adam et al. 2019). We more specifically address the following questions: - Can cemeteries play an important role in conserving plant biodiversity on a regional scale? And what are the specificities of the flora of cemeteries comparing with other urban areas? - What are the effect of urban-rural gradient on the flora of cemeteries? Material and methods The Île-de-France region and its cemeteries The climate is oceanic with continental trends; the mean annual temperature is about 11°C with an annual rainfall of about 650 mm (Météo France in Fernez et al. 2015). The land use pattern provided by the the National Institute of Statistics and Economic Studies (INSEE Dossier Île-de-France 2022) shows that artificial surfaces cover 23% of this region with a human density of about 1,022 inhabit./km 2 making it the most urbanised region in France. Agricultural land covers 50% of the region and forests and woodlands, 23% (Adam et al. 2019). Cemeteries have been inventoried throughout the Île-de-France region (Fig. 1). The cartography of the cemeteries comes from the "BD Topo" (Institut national de l’information géographique et forestière 2011). In this cartography, as some cemeteries are composed of several parts (military cemeteries vs. civil ones; Muslim and Jewish parts etc.), entities that are spatially less than 20 metres apart have been merged. Thus, there are 1,656 cemeteries in the Île-de-France region after this aggregation. Many contemporary French cemeteries were created in the 19th century (Bertrand 2015). Before the French Revolution, burial sites were scattered, narrow and embedded in the urban landcover, particularly around places of worship. A few of these cemeteries remain today around churches, often small in size, with a second, more recent cemetery outside the village centre. From the 19th century onwards (and up until the 1930s), burials were grouped together in large peri-urban cemeteries, for reasons of hygiene. During this period, cemeteries also became public places where people came to pay their respects or take a stroll; thus, the layout was rationalized with designated areas for burials, the delineation of main and secondary circulation paths, which could be lined with trees. A typical example of this type of layout is the famous Père-Lachaise cemetery in eastern Paris. In small villages, the rationalization of cemetery organization has sometimes been less strict, limited to delineating the cemetery with an enclosure wall and marking pathways. Pre-existing grassy areas may have been preserved instead of seeding a lawn; sometimes, graves were even directly installed within them. With urban sprawl, cemeteries that were initially established outside of cities and villages may now find themselves embedded within the urban matrix. Since several decades, some of the new French urban cemeteries have been the subject of renewed landscape research, often inspired by the park-like cemeteries of the Anglo-American world or Germanic Europe. Selection of the floristic dataset Floristic data were extracted from FLORA, the database of the National Botanical Conservatory of the Paris Basin (hereafter CBNBP), a French public organization aiming to study and protect the flora of this area. One of the aim of the botanists of the CBNBP, with the help both of competent amateurs and professionals, is to establish periodic floristic atlases to describe the geographical distribution of all species growing in the area. Since about 2000, they visited the ‘communes’ (French administrative municipalities), or 5 x 5 km squares for more recent periods, and made georeferenced relevés in the different habitats to record as many plant species as they could observe in the sampling unit. Within this framework, cemeteries are often inventoried as areas representing the flora of towns and villages. For this study, we intersected the floristic georeferenced relevés with the cemeteries of the layer of the "BD Topo"; relevés where 70% of the mapped area is included in the cemetery were selected. We filtered the relevés to include only those carried out after 2000, which represent the vast majority of the data extracted (99.4%). The mean area of the selected relevés is 6,086 m²± 452 se. Nomenclature follows the French Taxref version 14 (Gargominy et al. 2020). Taxa were generally processed according to the taxonomic rank of the species. However, in order to avoid bias relating to the uneven recording of critical segregates, apomictic taxa ( e.g. Rubus , Taraxacum , Hieracium , Rosa, etc.) were aggregated to the genus level or in aggregates; the same applies to taxa whose taxonomic rank has varied between versions of Taxref ( e.g. Arenaria serpyllifolia and A. leptoclados which were only subspecies in some Taxref versions). For convenience, we will still use the terms species throughout this paper. Data analyses This work focuses only on spontaneous vascular flora, i.e. plants that have not been planted or sown, particularly for ornamental purposes. A regional nativeness status was assigned to each species, based on the main regional status in the catalogue of the vascular flora of Île-de-France (Perriat et al. 2020). A few adaptations have been made when the species has a secondary status of regional nativeness; for example, when Myosotis sylvatica or Bupleurum rotundifolium are present in cemeteries, they are certainly subspontaneous ( i.e. planted species that only escape very locally) rather than native. Species are considered to be of conservation interest if they are included on at least one of the following lists: national or regional protection lists (respectively arrêté du 20 janvier 1982 2 and arrêté du 11 mars 1991 3 ), national (UICN France, FCBN, AFB & MNHN, 2018) and regional (Filoche 2014) IUCN Red Lists or listed as ZNIEFF species for Île-de-France (Filoche 2023). The ZNIEFF list is linked to a French policy (Lepareur et al. 2022) which aims to inventory all areas of high ecological value likely to be affected by development projects; lists of ZNIEFF species are regional and are generally based on Red Lists but excluding species linked to anthopogenic habitats and including others which are good indicators of natural habitats even if they are not threatened. In order to analyse the floristic pools of species from an ecological point of view, the species were assigned to major habitat types based on the regional work of Perriat (2018). Seven major habitat types were identified: Heaths and dry grasslands on acidic soil; Dry grasslands, heaths and herbaceous fringes on neutral or calcareous soils; Mesophilous meadows and grasslands; Herbaceous wet vegetation (amphibious communities; humid grasslands and tall herb communities; bogs and marsh; aquatic vegetation) Annual arable species; Forest (forest, edges and scrubs thickets) ; Anthropogenic vegetations (wastelands, weeds, wall vegetation). All data analyses, graphical representations and mapping were carried out using the software R (R Core Team 2022) with the following packages: ggplot2 (Wickham et al. 2021a), sf (Pebesma 2018), mapsf (Giraud 2022), plyr (Wickham 2011), dplyr (Wickham et al. 2021b) and indicspecies (De Cáceres and Legendre 2009). Comparison between the flora of cemeteries and those of other urbanised habitats Unlike other studies (Geoffroy et al. 2020; Itescu and Jeschke 2024) that compare the biodiversity of cemeteries with that of certain well-identified urban spaces such as parks, botanical gardens, and wastelands, we have taken a different approach. Since we have access to a significant database where surveys in urban contexts have been conducted in extremely varied urban spaces and interstices (those previously mentioned, as well as sidewalks, walls, vegetable gardens, lawns, flower beds, railways, road verges, urban forests etc.), we have chosen to compare the flora of cemeteries with urban flora defined as a whole regardless of the habitat. All relevés to which the botanist had assigned the CORINE BIOTOPES (European Commission: Directorate-General for Environment 1991) codes 85 (urban parks and large gardens) and 86 (towns, villages and industrial sites) were extracted from FLORA, the CBNBP database. Among these relevés, those already selected in the "cemetery" dataset were excluded. The same filters as those used for the "cemetery" dataset were applied to the nativeness of the species; the result was 4,609 relevés which contain a total of 185,391 records. A species indicator analysis (De Cáceres and Legendre 2009) between the relevés of "cemetery dataset" and those from of other urbanised habitats was carried out in order to highlight the species that were significantly more frequent and relatively exclusive to one of the two datasets. As the data available are presence-absence, the Phi coefficient of association was used with 999 permutations to test for significance. As recommended by De Cáceres, the Phi coefficient was corrected for the unequal number of records between the two datasets. The indicator species lists of the two groups were compared in terms of nativeness and habitat preferences using a χ² test or a Fisher test if the conditions for applying the χ² are not met (expected numbers too low). Effect of the urban-rural gradient on cemetery flora The built-up cover within a 500-metre radius of each cemetery was used to characterise the urban-rural context in which the cemetery was located; this measure of building density to characterise an urban-rural gradient is regularly used in ecology (Williams et al. 2015; Vallet et al. 2008). The built-up layer used comes from the two categories "undifferentiated buildings" and "industrial buildings" from the BD TOPO®/RGE produced by the Institut national de l'information géographique et forestière (2011). The urban-rural gradient associated with the cemeteries studied has the following characteristics (Fig. 1) : the median of built-up cover is 6%, the mean 12%, the minimum is 0 and the maximum 69%. To describe the response of species to the urban-rural gradient and identify which species are associated with cemeteries in an urban context (high percentage of built-up area) or in a rural context (low percentage of built-up area), the niche model approach proposed by Chisté et al. (2016) was used. This method has the advantage of being able to detect trends even for species that are infrequent. For each species, the average percentage of built-up area around the cemeteries where the species is present is calculated (observed abundance-weighted means (AWM)), followed by a randomization procedure, with an abundance of 1 for presence, 0 for absence. The proportion of simulated AWM values greater than the observed AWM value of the species (or less than) allows for tests of significance. Species with a significantly lower AWM value than the simulated values are associated with cemeteries with a low proportion of built-up area in their vicinity (thus, cemeteries in villages in a rural context), while species with a higher AWM value are, on the contrary, associated with cemeteries in cities in an urban context. The characteristics of the indicator species of the two groups will be compared using a χ² test or a Fisher's exact test. Results General characteristics of the dataset The selected dataset contains 66,973 records distributed in 3,365 relevés covering 817 cemeteries of the 1,656 occuring in Île-de-France (Fig. 1). The median value of the number of records per cemetery is 52 (mean = 82); eleven cemeteries have only one record, while the most well-inventoried has 2,506. Although it appears that cemeteries have been more intensively surveyed in Paris and its inner suburbs, the entire set of cemeteries in the region has been taken into account (Fig. 2). The inventories of the CBNBP dataset also cover the entire growing season (Online Resource 1). Therefore, we are able to have a very good general overview of the floristic pool of cemeteries at the regional scale with this dataset. In total, 731 spontaneous species have been observed in the inventoried cemeteries, of which 16% are naturalised. Among these naturalised species, eleven are listed as invasive non-native species in Île-de-France (Wegnez 2022) and are considered well-established in the region, including in natural environments (category of 'established invaders'). Twelve belong to the category of 'potential invaders,' meaning they are well-established but only in disturbed environments and currently do not pose a problem in terms of biodiversity impacts (see Online Resource 2 for list). Sixty-nine species have a status of conservation interest. Seven are protected (one nationally, the others regionally), 39 threatened according to the regional red list and 51 listed in the Île-de-France ZNIEFF species list (list in Online Resource 3). Two species ( Veronica triphyllos and Leonurus cardiaca ) are Near Threatened (NT) on the national Red List. Most of these species were observed in one or two cemeteries only but ten were inventoried in more than five cemeteries (up to thirteen cemeteries for Galium parisiense and Laphangium luteoalbum ). Two of these species of conservation interest, Veronica triphyllos and V. acinifolia are only known in the whole region from one cemetery (Ivry-sur-Seine, a municipality of Paris inner suburbs) (Conservatoires botaniques nationaux 2024). Fig. 3 shows a photographic pannel of the ten most frequent species of conservation interest. One-hundred and fourteen cemeteries host at least one species of conservation interest, i.e. 13% of the cemeteries inventoried. Seventy-three of these cemeteries host only one species of conservation interest. The map in Fig. 4 shows the number of species of conservation interest found per cemetery. Among the species of conservation interest, species of anthropogenic communities were well represented (n = 17) but the majority were linked to semi-natural habitats: dry grasslands, heaths and herbaceous fringes on neutral or calcareous soils (n = 22) and heaths and dry grasslands on acidic soil (n = 14) in particular (Fig. 5). As orchids have often been a focus of interest in studies on the flora of cemeteries (Löki et al. 2019a), we also made a small focus on their presence in the cemeteries of Île-de-France. Thirteen different orchid species were detected in 94 cemeteries ( i.e. 11.5% of the cemeteries studied). Five orchid species were present in only one cemetery, and four in more than 20 cemeteries (Table 1). Four orchids found in cemeteries are species of conservation interest in Île-de-France: Ophrys aranifera , Ophrys fuciflora , Orchis simia and Spiranthes spiralis . Table 1: List of the orchid species with the number of cemeteries where they were observed. Nom scientifique Nb cemeteries Himantoglossum hircinum (L.) Spreng., 1826 41 Ophrys apifera Huds., 1762 40 Anacamptis pyramidalis (L.) Rich., 1817 26 Epipactis helleborine (L.) Crantz, 1769 22 Ophrys aranifera Huds., 1778 6 Platanthera chlorantha (Custer) Rchb., 1828 6 Cephalanthera damasonium (Mill.) Druce, 1906 4 Neottia ovata (L.) Bluff & Fingerh., 1837 4 Anacamptis morio (L.) R.M.Bateman, Pridgeon & M.W.Chase, 1997 1 Ophrys fuciflora (F.W.Schmidt) Moench, 1802 1 Orchis purpurea Huds., 1762 1 Orchis simia Lam., 1779 1 Spiranthes spiralis (L.) Chevall., 1827 1 Comparison of the flora of cemeteries with the flora of other urban habitats The number of indicator species of cemeteries was 288, of which 26 are species of conservation interest. The number of indicator species in other urban habitats was 73, none of which are species of conservation interest. Indicator species lists are in Online Resource 4. The proportion of native and naturalized species between the two groups of indicator species did not differ significantly (χ² test, p-value = 0.45); around 80% of the indicator species in either context were native (Fig. 6). Both in cemeteries and in other urban habitats, a significant proportion of indicator species are linked to anthropogenic vegetation. Nevertheless, the proportion of indicator species between the seven habitat types is significantly different (Fisher test, p < 0,001) (Fig. 7). All species of heathland and dry neutrophilous or calcareous grasslands, as well as the vast majority (3 vs. 67) of species of mesophilous grasslands, are found only among the indicator species of cemeteries. Conversely, species associated with wet herbaceous vegetation and forest vegetation are overepresented among the indicator species of other urban areas. The effect of the urban-rural gradient on the flora of cemeteries The number of species whose distribution is significantly linked to cemeteries in an urban context is 148, of which three are of conservation interest. The number linked to cemeteries in a rural context is 130, of which 14 are of conservation interest. The detailed lists can be consulted in Online Resource 5. The vast majority of species whose distribution is linked to rural cemeteries are native (Fig. 8): only three are naturalized non-natives, whereas this proportion is 38% for species linked to cemeteries in an urban context (χ² test, p < 0.001). There are significant differences in the habitat preference of species associated with cemeteries in urban and rural environments (Fisher's exact test, p < 0.001) (Fig. 9). Species associated with heathland, and dry neutrophilous or calcareous grasslands and meadows are over-represented in species associated with rural cemeteries, whereas those associated with forests and margins and anthropogenic environments are much more common in species associated with urban cemeteries. Discussion Many studies on biodiversity in cemeteries have focused on cemeteries that are remarkable for their size, architecture or history (Buchholz et al. 2016 ; Buldrini et al. 2023 ; Kowarik et al. 2016 ; Otves, Arsene et al. 2016 ; Šilc 2009 ; Yılmaz et al. 2018 ). The present study focuses on a synthesis of the floristic biodiversity of cemeteries across an entire region with inventories available for half of the cemeteries in this region, i.e. 817 cemeteries. Contrary to many studies in which inventories are carried out on a single visit (Löki et al. 2019b ; Konic et al. 2021 ), the present dataset covers the entire vegetation period (April to October), which makes it possible to draw up a fairly exhaustive portrait of the flora that develops in the cemeteries of Île-de-France, whatever its phenology. In this way, vernal species such as those associated with dry grasslands on acidic sands, and naturalised species, many of which tend to appear late in the season, are not overlooked. Our results may be nuanced by the fact that our cemetery inventories are not necessarily limited to the strict perimeter of cemeteries, but may include some adjacent areas such as the foot of walls outside the cemetery enclosure, some adjacent streets or the cemetery car park. For this reason, our work concerns the floristic diversity of cemeteries and their adjacent areas. For a regional view of the biodiversity of cemeteries, the inclusion of these adjacent areas is probably negligible since it is unlikely that the species in the adjacent areas are specific to them. From an applied point of view, these adjacent areas are very often public, urbanised and managed by the same services as the cemetery, so it is not inconsistent for them to be included in this work. The importance of cemeteries for the conservation of plant biodiversity The 731 species observed in the 817 cemeteries surveyed in the Île-de-France region represent almost half of the spontaneous flora observed at the same time in the region (around 1,530 species, calculated according to the regional list (Perriat et al. 2020 )), a proportion comparable to that found to a study in the Rurh Metropolitan Region in Germany (Buch et al. 2025 ). Given the small area of cemeteries in the region, this figure is not negligible (Vallet 2024 ). The high species richness of cemeteries can probably be explained by the fact that there is an important diversity of micro-habitats in cemeteries: walls, very sunny areas and very shaded areas (for example, at the foot of the perimeter walls), grassy areas and, in some cemeteries, a few trees or shrubs, etc . In addition, the fact that there is a degree of disturbance (mechanical weeding, trampling, soil disturbance to create new graves, etc. ) can also encourage species diversity (Grime 1973 ; Nowińska et al. 2020 ). The proportion of non-native species in this cemetery flora pool (16%) is comparable to that of the regional flora (13.5%). This proportion is comparable to that observed in the cemetery of Modena (16.9%) (Buldrini et al. 2023 ) and much lower than that observed in Poland (33% on average for 78 cemeteries surveyed) (Nowińska et al. 2020 ). However, it is always complicated to analyse this type of figure, as it depends on the regional pool of species present, the climate, and a range of other factors like the city size (Pyšek 1998 ). Furthermore, in our study, we chose to exclude species that were not spontaneous, including subspontaneous species, i.e. planted species that only escape very locally. It is not always easy in the field to identify whether the species observed has been planted or not ( e.g. Sedum ); it is possible that we have been more restrictive in our study than in those previously cited. However, the place of non-native species also depends on the type of funerary practice (Buldrini et al. 2023 ; Nowińska et al. 2020 ; Gudžinskas 2005 ). In the Île-de-France region, the species planted to decorate graves are often poorly adapted to the local climate, as for example, All Saints' Day Chrysanthemums ( Chrysanthemum ×grandiflorum ), begonias as Begonia ×semperflorens-cultorum or horticultural cultivars that do not naturalise very well (la “pensée”, Viola ×wittrockiana ). Otherwise, more robust species are often planted in pots (Larramendy 2017 ) or in spaces with thin substrate they also die out during summer droughts. This probably limits the presence of non-native species from horticultural origin in our inventories, even if they are present ( Syringa vulgaris , Laburnum anagyroides , Robinia pseudoacacia , etc. ). These species are probably more related to the cemetery's ornamental green spaces than to plants introduced by private individuals to decorate graves. Less than 10% of the non-native species found in cemeteries are considered as invasive aliens in the Île-de-France region. The species present are considered to be well-established in the region; for these species, control measures are only recommended in special cases, particularly when these species invade natural habitat with conservation interest, and not in anthropogenic contexts (Wegnez 2022 ). In our study, of the 935 species present in cemeteries and/or other urban habitats (streets, wasteland, parks etc. ), 31% are significantly more frequent in cemeteries than other urban habitats. For species of conservation interest, this proportion is 51% (144 species of conservation interest recorded out of the 935 species in this dataset). These figures are quite considerable when compared with the small surface area of cemeteries in relation to other urban environments. Since the proportion of native and non-native species between those that prefer cemeteries and those that prefer other urban habitats does not differ, this may indicate that cemeteries are not a particularly important source for the introduction of non-native species, contrary to the conclusion of Walusiak and Krztoń ( 2021 ) in Poland. Nevertheless, of the 288 species found preferentially in cemeteries, 40 are non-native, three of which are considered invasive aliens ( Acer negundo , Ailanthus altissima and Parthenocissus inserta ); these species could be introduced or find a favourable habitat in cemeteries although they are not unique locations in that respect. Cemeteries are unique in that they provide habitats for species of dry or mesophilous herbaceous environments (dry grasslands on acid and calcareous soils (Fig. 5 ), meadows); conversely, species of woodland or wetland habitats are rarely found in cemeteries. In these urban contexts, these woodland and wetland species are probably associated with urban parks or riverbanks. With the exception of park-like cemeteries, these habitats are not very common in cemeteries. The fact that the flora of cemeteries is linked to dry habitats is not surprising (Buldrini et al. 2023 ; Buch et al. 2025 ). What is notable is that cemeteries are home to a pool of species associated with oligotrophic habitats, with both acidic and basic pH levels; these ecological traits are particularly rare among plant species in an urban context (Williams et al. 2015 ). It is therefore important to conserve these species and in particular the semi-natural remnant habitats that may exist in some cemeteries, and are of great interest for the conservation of biodiversity in an anthropogenic environment (Löki et al. 2020 ; Buch et al. 2025 ). Conservation issues differ between town and village cemeteries Although cemeteries are generally located in urbanised areas, the Île-de-France region is a good case of study for assessing the influence of an urban-rural gradient, since there is a highly contrasting gradient in the proportion of built-up areas around cemeteries, from cemeteries with less than 1% of built-up areas to cemeteries with 70% of built-up areas. The former probably correspond to village cemeteries, while those with the most built-up areas are cemeteries in large towns. We find a classic pattern in urban ecology (Hill et al. 2002 ), namely that the city favours the presence of non-native species. It can be seen that the majority of species whose distribution is linked to cemeteries with a highly built-up landscape are non-native species, whereas species linked to low percentages of built-up areas are almost all native. This result does not mean that non-native species are absent from rural cemeteries, but that the non-native species they have are present in cemeteries regardless of the level of urbanisation; in other words, it means that cemeteries in highly urbanised contexts have a pool of non-native species that are not found in cemeteries in villages. This means that it is the landscape context that seems to favour the presence of non-native species in cemeteries, rather than the cemeteries themselves being major points of introduction for non-native species. There are also differences in the habitat preferences of species associated with cemeteries in towns and villages. For example, the species assemblages of dry or mesophilous herbaceous environments (acid and limestone grasslands, meadows, heathlands), which have already been mentioned because some of their species are of conservation interest, are more closely associated with rural cemeteries. There are two possible reasons for this (Trzaskowska and Karczmarz 2013 ): urban cemeteries were probably established in areas yet influenced by human activities at the urban fringe with few remnant natural habitats. Morevore, the organization of urban cemeteries was more often planified by landscape architects with little space to pre-existing spontaneous vegetation whereas in villages pre-existing vegetation has sometimes been conserved (Bertrand 2015 ). The species that develop preferentially in urban environments tend to be associated with anthropogenic vegetation or forest vegetation. In the case of the latter, this could be linked to the recent development of so-called park-like cemeteries in towns; with vast areas of urban lawns and wooded parts, their flora is more similar to that of urban parks. Conclusion about cemeteries management practices Cemeteries play a particularly important role in preserving plant biodiversity in an anthropogenic context. In terms of conservation, particular attention must be paid to cemeteries, which contain areas of semi-natural remnant habitats; it must be a priority to protect these habitats and their associated species. In relation to this, it might be useful to identify cemeteries with remnant natural habitats to bring these conservation issues to the attention of local authorities. These semi-natural habitats should not be destroyed when the areas reserved for graves are enlarged. Their management should also be adapted: no fertilisation and less frequent mowing, as in many cases these areas are managed as lawns. Annual mowing at the end of the season, if possible with arisings removed, would be sufficient. As these areas are generally located on oligotrophic soils, the development of vegetation is limited. In addition, many of these areas have species that flower quite brightly ( Calluna vugaris and Erica cinerea, Scabiosa columbaria , Knautia arvensis, Armeria arenaria etc.), which are very interesting from an ornamental point of view. So local managers of cemeteries should be informed of these conservation issues and advised on the type of management to be implemented. Moreover, education of cemetery users to support the acceptance of less intensive management, leading to the presence of spontaneous flora in paths (Fig. 5 F) and the presence of tall (but flowering) vegetation in grassy areas would be valuable. In addition to the value of these semi-natural habitats, cemeteries host a range of diverse species that are not often found in other habitats of towns and villages. We encourage the maintenance of the diversity of habitats found in cemeteries for example by keeping old walls and maintaining both sunny and shaded areas. Maintaining the diversity of geological and soil contexts is also a factor of diversity on a regional scale (Buch et al. 2025 ); when developing cemeteries, local substrates should be favoured over importation of topsoil from outside. However, some recent practices linked to the prohibition of the use of pesticides have raised questions about whether the originality of cemetery floras can be maintained. Park-like cemeteries may host a large number of species, but the species found there are largely the same as those found in parks (species of urban lawns and wooded areas). So, on a city scale, their contribution can be rather redundant with parks already present. New pathway management practices that are developing as a result of the stopping of the use of pesticides raise questions about maintaining the specific characteristics of cemetery floras. The waterproofing of pathways (bitumen, cement) greatly reduces the surface area of favourable habitat; this should be limited to certain main pathways at most, leaving at least the secondary pathways permeable. The transformation of pathways in lawns by sowing of grasses (often dominated by red fescues Festuca ) also raises questions. A number of species characteristic of cemeteries are linked to mineral pathways (sand, alluvium), which provide them conditions of sunshine, dryness and low competition (Buch et al. 2025 ) that they will not find in pathways with dense grass cover. These may be, for example, species associated with dry sandy grasslands or with crops, such as different species of poppies ( Papaver rhoeas, P. dubium, P. argemone ) (Fig. 5 ) or field marigold ( Calendula arvensis ), or species associated with disturbed environments, such as Galium parisiense . It therefore seems important to retain mineral pathways and to allow them to be grassed over naturally (without sowing) in order to maintain the originality of the floral communities in cemeteries, whether they are linked to « ordinary » or conservation interest flora. We therefore strongly encourage the maintenance of paths that are neither waterproofed nor grassed over, with mechanical weed management. Abbreviations follows the French Taxref version 14 (Gargominy et al. 2020). Taxa were generally processed according to the taxonomic rank of the species. However in order to avoid bias relating to the uneven recording of critical segregates,apomictic taxa ( e.g. Rubus , Taraxacum , Hieracium , Rosa ,etc.) were aggregated to the genus level or in aggregates the same applies to taxa whose taxonomic rank has varied between versions of Taxref ( e.g. Arenaria serpyllifolia and A. leptoclados which were only subspecies in some Taxref versions). For convenience we will still use the terms species throughout this paper. Declarations Funding This study was initiated thanks to the financial support of "L'Institut Paris Région – Agence régionale de la biodiversité (ARB)". Competing Interests The author declare she has any interest in the current study. Data Availability The datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request. Author Contribution J.V. designed the study, performed the statistical analyses and wrote the manuscript. Acknowledgement I would like to sincerely thank Oliver Pescott and Hervé Daniel for their encouragement in publishing this work and for their advice opinion on the first draft of this paper References Adam M, Cormier T and Gobled L (2019). Vers un développement urbain raisonné et durable. Note Rapide IAU n°797. https://institutparisregion.fr/fileadmin/NewEtudes/Etude_1762/NR_797_web.pdf. 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Supplementary Files supp.docx Cite Share Download PDF Status: Published Journal Publication published 09 Apr, 2026 Read the published version in Urban Ecosystems → Version 1 posted Editorial decision: Revision requested 27 Oct, 2025 Reviews received at journal 17 Oct, 2025 Reviews received at journal 01 Oct, 2025 Reviewers agreed at journal 04 Sep, 2025 Reviewers agreed at journal 03 Sep, 2025 Reviewers invited by journal 02 Sep, 2025 Editor assigned by journal 27 Aug, 2025 Submission checks completed at journal 27 Aug, 2025 First submitted to journal 24 Aug, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7448159","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":510251812,"identity":"6b7c8c4c-c629-4ca1-b804-4b31f2ac0cf8","order_by":0,"name":"Jeanne Vallet","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6ElEQVRIie3PMQuCQBTA8fdwaLFdOLCvYAhF0IdJWiOCIBwaTg50EVpvkPoKTc4dgi5+gKAphFpriaboNBpaDseG+08PHj/eHYBO94ch/U4dQ44+gCPnWztiyPFQNgR5u3s1EWELYkTsUi188DYMg+q+TedDQpH5qofF+dDlJXg8Q+aI9LQcJQcMShXhkwHphuBChqElibc/TjpnqiK764N0X+D2MoyeImkIBkrCTXmFgu3IKyBoGxLPVsTMLbsv/2KVeU28QEn6UZEScz027YKJm7+uyVSoyWdp/VxWAYCecqvT6XS6ujcD01RusHb4SgAAAABJRU5ErkJggg==","orcid":"","institution":"Muséum national d'histoire naturelle","correspondingAuthor":true,"prefix":"","firstName":"Jeanne","middleName":"","lastName":"Vallet","suffix":""}],"badges":[],"createdAt":"2025-08-24 20:23:08","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7448159/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7448159/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11252-026-01939-z","type":"published","date":"2026-04-09T15:57:57+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":90913610,"identity":"fc9c5f96-15cd-4e42-a2fb-d0a3c1ee0e3c","added_by":"auto","created_at":"2025-09-09 13:53:58","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":362301,"visible":true,"origin":"","legend":"\u003cp\u003ea. Location of the Île-de-France region in France with its capital Paris. b. Location of inventoried (pink circles symbols) and non-inventoried cemeteries (black squares symbols) in Île-de-France. The size of the pink circle is proportional to the percentage of built-up area within a 500-metre radius of each cemetery. The main rivers are shown in blue and the limits of départements in black\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/0a3013290e7332e350661bce.png"},{"id":90909316,"identity":"69c1a089-47bb-4df9-b6fb-b23c2f43b710","added_by":"auto","created_at":"2025-09-09 13:29:48","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":283723,"visible":true,"origin":"","legend":"\u003cp\u003eMaps of inventoried cemeteries after 2000 in Île-de-France. The size of circles is proportional to: a. Number of records per cemetery. b. Species richness per cemetery\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/dc96d856f24e04c11024615b.png"},{"id":90909432,"identity":"f9a9aec9-0eb5-4be9-963c-ed359a205135","added_by":"auto","created_at":"2025-09-09 13:30:09","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1246664,"visible":true,"origin":"","legend":"\u003cp\u003eIllustration of the ten most frequent species of conservation interest; we precise the number of cemeteries in which they were found and their habitat and their main habitat\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/f42a06b0682042f9dd56e580.png"},{"id":90913608,"identity":"84b32621-00db-45c9-a5a2-817ba991bc8b","added_by":"auto","created_at":"2025-09-09 13:53:57","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":261833,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the number of species of conservation interest inventoried per cemetery (post-2000 observations). The size of circles is proportional to the number of species of conservation interest; orange triangles locate the cemeteries surveyed where no species of conservation interest were found\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/d7611f728978f0f6d46cf8c8.png"},{"id":90909320,"identity":"fa058408-7ee6-486c-9d03-7a63a9bcaf6c","added_by":"auto","created_at":"2025-09-09 13:29:48","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":1332544,"visible":true,"origin":"","legend":"\u003cp\u003eIllustration of the vegetation in different cemeteries in Île-de-France.; a. An area of dry calcareous grassland in cemetery of Ury (© J. Wegnez, CBNBP/MNHN); b. Acidic sandy grassland in a path in the cemetery of Nonville; c. Mesophilic grassland in the cemetery of les Marêts (© J. Vallet, CBNBP/MNHN); d. Heathland vegetation in the cemetery of Janvry (©T. Fernez, CBNBP/MNHN); e. A poppy (\u003cem\u003ePapaver argemone\u003c/em\u003e), an annual arable species in the sandy paths of a cemetery (© J. Vallet, CBNBP/MNHN); f. Mineral path with spontaneous development of vegetation and ornamental plants on graves (partially died because of the drought) at La Garenne-Colombes (© J. Vallet, CBNBP/MNHN)\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/fc5f978a3589821a593477b6.png"},{"id":90912021,"identity":"ff1ac87c-4f5b-4d5c-aefd-54adf3b644e3","added_by":"auto","created_at":"2025-09-09 13:45:48","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":45725,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of species number of indicator species of cemeteries and other urban habitats according to their nativeness. The width of each column is proportional to the number of indicator species in each group (preference for \"cemeteries\" or \"other urbanised habitats\") and the height of each rectangle proportional to the number of species by nativeness category\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/9c5fc5eeb96163876dec7f7a.png"},{"id":90910360,"identity":"5765e1d7-1149-4152-bef7-3d633be7da49","added_by":"auto","created_at":"2025-09-09 13:37:48","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":113446,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of species number of indicator species of cemeteries and other urban habitats according to their habitat type. The width of each column is proportional to the number of indicator species in each group (preference for \"cemeteries\" or \"other urbanised habitats\") and the height of each rectangle proportional to the number of species by habitat types\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/5b844d04e9acac7079db75c7.png"},{"id":90912024,"identity":"c731dbe8-67bd-4d14-a749-0834cbddeaf7","added_by":"auto","created_at":"2025-09-09 13:45:48","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":34411,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of species number of indicator species of urban and rural cemeteries according to their nativeness. The width of each column is proportional to the number of indicator species in each group (preference for cemeteries in urban or rural contexts) and the height of each rectangle proportional to the number of species by nativeness category\u003c/p\u003e","description":"","filename":"8.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/09994165c03c8c756296f597.png"},{"id":90910357,"identity":"58c48a30-9113-4b4d-86c8-9841dc141c01","added_by":"auto","created_at":"2025-09-09 13:37:48","extension":"png","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":76000,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of species number of indicator species of urban and rural cemeteries according to their habitat type. The width of each column is proportional to the number of indicator species in each group (preference for cemeteries in urban or rural contexts) and the height of each rectangle proportional to the number of species by habitat types\u003c/p\u003e","description":"","filename":"9.png","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/9501214547a1e1472c143dec.png"},{"id":106809850,"identity":"d04df009-3b43-41e4-a29d-387cf3b4c27a","added_by":"auto","created_at":"2026-04-13 16:13:11","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4113384,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/968c889e-b86a-47ac-8157-e10c70db9fc1.pdf"},{"id":90910353,"identity":"e2d4737f-6b9f-4b3e-9b7c-4921d4f37c8b","added_by":"auto","created_at":"2025-09-09 13:37:48","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":70462,"visible":true,"origin":"","legend":"","description":"","filename":"supp.docx","url":"https://assets-eu.researchsquare.com/files/rs-7448159/v1/22bcc8bc7734ddf9e0e40389.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"What role can cemeteries play in conserving plant biodiversity in different urbanisation contexts?","fulltext":[{"header":"Introduction","content":"\u003cp\u003eGiven the extent of land use transformations on a planetary scale due to the development of urbanization and agriculture, it is now essential to also consider the conservation of biodiversity in these anthropogenic spaces (Planchuelo et al. 2019). Different approaches can be envisaged to take better account of biodiversity in these anthropogenic contexts: Planchuelo et al (2019) stress the importance of all types of ecosystem for the conservation of plant species of conservation interest, whether they have been profoundly altered by human activities (novel ecosystems) or are made up of fragments of relict habitats (remnant ecosystems) or a hybrid between the two.\u003c/p\u003e\n\u003cp\u003eCemeteries are found in almost every town and village; they can vary greatly in terms of size, age, layout, funeral practices and the degree of urbanisation in the surrounding area. Generally speaking, cemeteries fall into the category of new ecosystems (Planchuelo et al. 2019), but some cemeteries have the particularity of harbouring remant habitats that may be of great interest for the conservation of plant species (L\u0026ouml;ki et al. 2020; Moln\u0026aacute;r et al. 2017). Cemeteries have also been identified as a potential source of non-native species, some of which may be invasive (Walusiak and Krztoń 2021). Many studies on cemetery flora have focused on a few cemeteries that are often singular in terms of their characteristics (Buldrini et al. 2023; Yılmaz et al. 2018; Yılmaz et al. 2018) and few have taken a more global approach on the scale of an entire territory (Buch et al. 2025). Moreover, as funerary practices can vary widely between countries, religions etc., the results cannot necessarily be more broadly generalised. Few studies have been conducted on the biodiversity of cemeteries in Western Europe (Itescu and Jeschke 2024). Finally, the 2020s mark a pivotal period in terms of management practices for cemeteries in France (European Environment Agency 2023) because since the 1\u003csup\u003est\u003c/sup\u003e of July 2022, pesticides are prohibited in these areas\u003ca href=\"#_ftn1\" name=\"_ftnref1\" title=\"\"\u003e\u003c/a\u003e\u003csup\u003e1\u003c/sup\u003e; a number of municipalities have anticipated the law (Flandin 2019). It is therefore particularly important to identify the state of present biodiversity in cemeteries in order to understand the future effects of this change in management.\u003c/p\u003e\n\u003cp\u003eThe aim of this study is to examine the vascular plant biodiversity of cemeteries in \u0026Icirc;le-de-France, a region surrounding Paris where human pressure is particularly strong, with agricultural areas occupying around 50% of the regional surface area and urban areas 23% (Adam et al. 2019).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWe more specifically address the following questions:\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e- Can cemeteries play an important role in conserving plant biodiversity on a regional scale? And what are the specificities of the flora of cemeteries comparing with other urban areas?\u003c/p\u003e\n\u003cp\u003e- What are the effect of urban-rural gradient on the flora of cemeteries?\u0026nbsp;\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cp\u003e\u003cem\u003eThe \u0026Icirc;le-de-France region and its cemeteries\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe climate is oceanic with continental trends; the mean annual temperature is about 11\u0026deg;C with an annual rainfall of about 650 mm (M\u0026eacute;t\u0026eacute;o France \u003cem\u003ein\u0026nbsp;\u003c/em\u003eFernez et al. 2015). The land use pattern provided by the the National Institute of Statistics and Economic Studies (INSEE Dossier \u0026Icirc;le-de-France 2022) shows that artificial surfaces cover 23% of this region with a human density of about 1,022 inhabit./km\u003csup\u003e2\u0026nbsp;\u003c/sup\u003emaking it the most urbanised region in France. Agricultural land covers 50% of the region and forests and woodlands, 23% (Adam et al. 2019).\u003c/p\u003e\n\u003cp\u003eCemeteries have been inventoried throughout the \u0026Icirc;le-de-France region (Fig. 1). The cartography of the cemeteries comes from the \u0026quot;BD Topo\u0026quot; (Institut national de l\u0026rsquo;information g\u0026eacute;ographique et foresti\u0026egrave;re 2011). In this cartography, as some cemeteries are composed of several parts (military cemeteries \u003cem\u003evs.\u0026nbsp;\u003c/em\u003ecivil ones;\u003cem\u003e\u0026nbsp;\u003c/em\u003eMuslim and Jewish parts etc.), entities that are spatially less than 20 metres apart have been merged. Thus, there are 1,656 cemeteries in the \u0026Icirc;le-de-France region after this aggregation.\u003c/p\u003e\n\u003cp\u003eMany contemporary French cemeteries were created in the 19th century (Bertrand 2015). Before the French Revolution, burial sites were scattered, narrow and embedded in the urban landcover, particularly around places of worship. A few of these cemeteries remain today around churches, often small in size, with a second, more recent cemetery outside the village centre. From the 19th century onwards (and up until the 1930s), burials were grouped together in large peri-urban cemeteries, for reasons of hygiene. During this period, cemeteries also became public places where people came to pay their respects or take a stroll; thus, the layout was rationalized with designated areas for burials, the delineation of main and secondary circulation paths, which could be lined with trees. A typical example of this type of layout is the famous P\u0026egrave;re-Lachaise cemetery in eastern Paris. In small villages, the rationalization of cemetery organization has sometimes been less strict, limited to delineating the cemetery with an enclosure wall and marking pathways. Pre-existing grassy areas may have been preserved instead of seeding a lawn; sometimes, graves were even directly installed within them. With urban sprawl, cemeteries that were initially established outside of cities and villages may now find themselves embedded within the urban matrix. Since several decades, some of the new French urban cemeteries have been the subject of renewed landscape research, often inspired by the park-like cemeteries of the Anglo-American world or Germanic Europe.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSelection of the floristic dataset\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eFloristic data were extracted from FLORA, the database of the National Botanical Conservatory of the Paris Basin (hereafter CBNBP), a French public organization aiming to study and protect the flora of this area.\u0026nbsp;One of the aim of the botanists of the CBNBP, with the help both of competent amateurs and professionals, is to establish periodic floristic atlases to describe the geographical distribution of all species growing in the area. Since about 2000, they visited the \u0026lsquo;communes\u0026rsquo; (French administrative municipalities), or 5\u0026nbsp;x 5 km squares for more recent periods, and made georeferenced relev\u0026eacute;s in the different habitats to record as many plant species as they could observe in the sampling unit. Within this framework, cemeteries are often inventoried as areas representing the flora of towns and villages. For this study, we intersected the floristic georeferenced relev\u0026eacute;s with the cemeteries of the layer of the \u0026quot;BD Topo\u0026quot;; relev\u0026eacute;s where 70% of the mapped area is included in the cemetery were selected. We filtered the relev\u0026eacute;s to include only those carried out after 2000, which represent the vast majority of the data extracted (99.4%). The mean area of the selected relev\u0026eacute;s is 6,086 m\u0026sup2;\u0026plusmn; 452 se.\u003c/p\u003e\n\u003cp\u003eNomenclature follows the French Taxref version 14 (Gargominy et al. 2020). Taxa were generally processed according to the taxonomic rank of the species. However, in order to avoid bias relating to the uneven recording of critical segregates, apomictic taxa (\u003cem\u003ee.g.\u003c/em\u003e \u003cem\u003eRubus\u003c/em\u003e, \u003cem\u003eTaraxacum\u003c/em\u003e, \u003cem\u003eHieracium\u003c/em\u003e, \u003cem\u003eRosa,\u0026nbsp;\u003c/em\u003eetc.) were aggregated to the genus level or in aggregates; the same applies to taxa whose taxonomic rank has varied between versions of Taxref (\u003cem\u003ee.g.\u003c/em\u003e \u003cem\u003eArenaria serpyllifolia\u003c/em\u003e and \u003cem\u003eA. leptoclados\u0026nbsp;\u003c/em\u003ewhich were only subspecies in some Taxref versions). For convenience, we will still use the terms species throughout this paper.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eData analyses\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThis work focuses only on spontaneous vascular flora, \u003cem\u003ei.e.\u003c/em\u003e plants that have not been planted or sown, particularly for ornamental purposes. A regional nativeness status was assigned to each species, based on the main regional status in the catalogue of the vascular flora of \u0026Icirc;le-de-France (Perriat et al. 2020). A few adaptations have been made when the species has a secondary status of regional nativeness; for example, when \u003cem\u003eMyosotis sylvatica\u003c/em\u003e or \u003cem\u003eBupleurum rotundifolium\u003c/em\u003e are present in cemeteries, they are certainly subspontaneous (\u003cem\u003ei.e.\u003c/em\u003e planted species that only escape very locally) rather than native.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSpecies are considered to be of conservation interest if they are included on at least one of the following lists: national or regional protection lists (respectively arr\u0026ecirc;t\u0026eacute; du 20 janvier 1982\u003ca href=\"#_ftn1\" name=\"_ftnref1\" title=\"\"\u003e\u003c/a\u003e\u003csup\u003e2\u003c/sup\u003e and arr\u0026ecirc;t\u0026eacute; du 11 mars 1991\u003ca href=\"#_ftn2\" name=\"_ftnref2\" title=\"\"\u003e\u003c/a\u003e\u003csup\u003e3\u003c/sup\u003e), national (UICN France, FCBN, AFB \u0026amp; MNHN, 2018) and regional (Filoche 2014) IUCN Red Lists or listed as ZNIEFF species for \u0026Icirc;le-de-France (Filoche 2023). The ZNIEFF list is linked to a French policy (Lepareur et al.\u003cem\u003e\u0026nbsp;\u003c/em\u003e2022) which aims to inventory all areas of high ecological value likely to be affected by development projects; lists of ZNIEFF species are regional and are generally based on Red Lists but excluding species linked to anthopogenic habitats and including others which are good indicators of natural habitats even if they are not threatened.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn order to analyse the floristic pools of species from an ecological point of view, the species were assigned to major habitat types based on the regional work of Perriat (2018). Seven major habitat types were identified:\u003c/p\u003e\n\u003cul class=\"decimal_type\"\u003e\n \u003cli\u003eHeaths and dry grasslands on acidic soil;\u003c/li\u003e\n \u003cli\u003eDry grasslands, heaths and herbaceous fringes on neutral or calcareous soils;\u003c/li\u003e\n \u003cli\u003eMesophilous meadows and grasslands;\u003c/li\u003e\n \u003cli\u003eHerbaceous wet vegetation (amphibious communities; humid grasslands and tall herb communities; bogs and marsh; aquatic vegetation)\u003c/li\u003e\n \u003cli\u003eAnnual arable species;\u003c/li\u003e\n \u003cli\u003eForest (forest, edges and scrubs thickets) ;\u003c/li\u003e\n \u003cli\u003eAnthropogenic vegetations (wastelands, weeds, wall vegetation).\u003c/li\u003e\n\u003c/ul\u003e\n\u003cp\u003eAll data analyses, graphical representations and mapping were carried out using the software R (R Core Team 2022) with the following packages: ggplot2 (Wickham et al. 2021a), sf (Pebesma 2018), mapsf (Giraud 2022), plyr (Wickham 2011), dplyr (Wickham et al.\u003cem\u003e\u0026nbsp;\u003c/em\u003e2021b) and indicspecies (De C\u0026aacute;ceres and Legendre 2009).\u003c/p\u003e\n\u003cp\u003e\u003cspan id=\"_Toc152696761\"\u003e\u003cem\u003eComparison between the flora of cemeteries and those of other urbanised habitats\u0026nbsp;\u003c/em\u003e\u003c/span\u003e\u003c/p\u003e\n\u003cp\u003eUnlike other studies (Geoffroy et al. 2020; Itescu and Jeschke 2024) that compare the biodiversity of cemeteries with that of certain well-identified urban spaces such as parks, botanical gardens, and wastelands, we have taken a different approach. Since we have access to a significant database where surveys in urban contexts have been conducted in extremely varied urban spaces and interstices (those previously mentioned, as well as sidewalks, walls, vegetable gardens, lawns, flower beds, railways, road verges, urban forests etc.), we have chosen to compare the flora of cemeteries with urban flora defined as a whole regardless of the habitat. All relev\u0026eacute;s to which the botanist had assigned the CORINE BIOTOPES (European Commission: Directorate-General for Environment 1991) codes 85 (urban parks and large gardens) and 86 (towns, villages and industrial sites) were extracted from FLORA, the CBNBP database. Among these relev\u0026eacute;s, those already selected in the \u0026quot;cemetery\u0026quot; dataset were excluded. The same filters as those used for the \u0026quot;cemetery\u0026quot; dataset were applied to the nativeness of the species; the result was 4,609 relev\u0026eacute;s which contain a total of 185,391 records.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eA species indicator analysis (De C\u0026aacute;ceres and Legendre 2009) between the relev\u0026eacute;s of \u0026quot;cemetery dataset\u0026quot; and those from of other urbanised habitats was carried out in order to highlight the species that were significantly more frequent and relatively exclusive to one of the two datasets. As the data available are presence-absence, the Phi coefficient of association was used with 999 permutations to test for significance. As recommended by De C\u0026aacute;ceres, the Phi coefficient was corrected for the unequal number of records between the two datasets. The indicator species lists of the two groups were compared in terms of nativeness and habitat preferences using a \u0026chi;\u0026sup2; test or a Fisher test if the conditions for applying the \u0026chi;\u0026sup2; are not met (expected numbers too low). \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eEffect of the urban-rural gradient on cemetery flora\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe built-up cover within a 500-metre radius of each cemetery was used to characterise the urban-rural context in which the cemetery was located; this measure of building density to characterise an urban-rural gradient is regularly used in ecology (Williams et al. 2015; Vallet et al. 2008). The built-up layer used comes from the two categories \u0026quot;undifferentiated buildings\u0026quot; and \u0026quot;industrial buildings\u0026quot; from the BD TOPO\u0026reg;/RGE produced by the Institut national de l\u0026apos;information g\u0026eacute;ographique et foresti\u0026egrave;re (2011). The urban-rural gradient associated with the cemeteries studied has the following characteristics (Fig. 1) : the median of built-up cover is 6%, the mean 12%, the minimum is 0 and the maximum 69%.\u003c/p\u003e\n\u003cp\u003eTo describe the response of species to the urban-rural gradient and identify which species are associated with cemeteries in an urban context (high percentage of built-up area) or in a rural context (low percentage of built-up area), the niche model approach proposed by Chist\u0026eacute; et al. (2016) was used. This method has the advantage of being able to detect trends even for species that are infrequent. For each species, the average percentage of built-up area around the cemeteries where the species is present is calculated (observed abundance-weighted means (AWM)), followed by a randomization procedure, with an abundance of 1 for presence, 0 for absence. The proportion of simulated AWM values greater than the observed AWM value of the species (or less than) allows for tests of significance. Species with a significantly lower AWM value than the simulated values are associated with cemeteries with a low proportion of built-up area in their vicinity (thus, cemeteries in villages in a rural context), while species with a higher AWM value are, on the contrary, associated with cemeteries in cities in an urban context. The characteristics of the indicator species of the two groups will be compared using a \u0026chi;\u0026sup2; test or a Fisher\u0026apos;s exact test.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003eGeneral characteristics of the dataset\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe selected dataset contains 66,973 records distributed in 3,365 relev\u0026eacute;s covering 817 cemeteries of the 1,656 occuring in \u0026Icirc;le-de-France (Fig. 1). The median value of the number of records per cemetery is 52 (mean = 82); eleven cemeteries have only one record, while the most well-inventoried has 2,506. Although it appears that cemeteries have been more intensively surveyed in Paris and its inner suburbs, the entire set of cemeteries in the region has been taken into account (Fig. 2). The inventories of the CBNBP dataset also cover the entire growing season (Online Resource 1). Therefore, we are able to have a very good general overview of the floristic pool of cemeteries at the regional scale with this dataset.\u003c/p\u003e\n\u003cp\u003eIn total, 731 spontaneous species have been observed in the inventoried cemeteries, of which 16% are naturalised. Among these naturalised species, eleven are listed as invasive non-native species in \u0026Icirc;le-de-France (Wegnez 2022) and are considered well-established in the region, including in natural environments (category of \u0026apos;established invaders\u0026apos;). Twelve belong to the category of \u0026apos;potential invaders,\u0026apos; meaning they are well-established but only in disturbed environments and currently do not pose a problem in terms of biodiversity impacts (see Online Resource 2 for list).\u003c/p\u003e\n\u003cp\u003eSixty-nine species have a status of conservation interest. Seven are protected (one nationally, the others regionally), 39 threatened according to the regional red list and 51 listed in the \u0026Icirc;le-de-France ZNIEFF species list (list in Online Resource 3). Two species (\u003cem\u003eVeronica triphyllos\u003c/em\u003e and \u003cem\u003eLeonurus cardiaca\u003c/em\u003e) are Near Threatened (NT) on the national Red List. Most of these species were observed in one or two cemeteries only but ten were inventoried in more than five cemeteries (up to thirteen cemeteries for \u003cem\u003eGalium parisiense\u003c/em\u003e and \u003cem\u003eLaphangium luteoalbum\u003c/em\u003e). Two of these species of conservation interest, \u003cem\u003eVeronica triphyllos\u0026nbsp;\u003c/em\u003eand \u003cem\u003eV. acinifolia\u0026nbsp;\u003c/em\u003eare only known in the whole region from one cemetery (Ivry-sur-Seine, a municipality of Paris inner suburbs) (Conservatoires botaniques nationaux 2024). Fig. 3 shows a photographic pannel of the ten most frequent species of conservation interest. One-hundred and fourteen cemeteries host at least one species of conservation interest, \u003cem\u003ei.e.\u003c/em\u003e 13% of the cemeteries inventoried. Seventy-three of these cemeteries host only one species of conservation interest. The map in Fig. 4 shows the number of species of conservation interest found per cemetery. Among the species of conservation interest, species of anthropogenic communities were well represented (n = 17) but the majority were linked to semi-natural habitats: dry grasslands, heaths and herbaceous fringes on neutral or calcareous soils (n = 22) and heaths and dry grasslands on acidic soil (n = 14) in particular (Fig. 5).\u003c/p\u003e\n\u003cp\u003eAs orchids have often been a focus of interest in studies on the flora of cemeteries (L\u0026ouml;ki et al. 2019a), we also made a small focus on their presence in the cemeteries of \u0026Icirc;le-de-France. Thirteen different orchid species were detected in 94 cemeteries (\u003cem\u003ei.e.\u0026nbsp;\u003c/em\u003e11.5% of the cemeteries studied). Five orchid species were present in only one cemetery, and four in more than 20 cemeteries (Table 1). Four orchids found in cemeteries are species of conservation interest in \u0026Icirc;le-de-France: \u003cem\u003eOphrys aranifera\u003c/em\u003e, \u003cem\u003eOphrys fuciflora\u003c/em\u003e, \u003cem\u003eOrchis simia\u003c/em\u003e and \u003cem\u003eSpiranthes spiralis\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 1:\u003c/strong\u003e List of the orchid species with the number of cemeteries where they were observed.\u003c/p\u003e\n\u003cdiv align=\"\"\u003e\n \u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"502\"\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNom scientifique\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNb cemeteries\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eHimantoglossum hircinum\u003c/em\u003e (L.) Spreng., 1826\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e41\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eOphrys apifera\u003c/em\u003e Huds., 1762\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e40\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eAnacamptis pyramidalis\u003c/em\u003e (L.) Rich., 1817\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e26\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eEpipactis helleborine\u003c/em\u003e (L.) Crantz, 1769\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eOphrys aranifera\u003c/em\u003e Huds., 1778\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003ePlatanthera chlorantha\u003c/em\u003e (Custer) Rchb., 1828\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eCephalanthera damasonium\u003c/em\u003e (Mill.) Druce, 1906\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eNeottia ovata\u003c/em\u003e (L.) Bluff \u0026amp; Fingerh., 1837\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eAnacamptis morio\u003c/em\u003e (L.) R.M.Bateman, Pridgeon \u0026amp; M.W.Chase, 1997\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eOphrys fuciflora\u003c/em\u003e (F.W.Schmidt) Moench, 1802\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eOrchis purpurea\u003c/em\u003e Huds., 1762\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eOrchis simia\u003c/em\u003e Lam., 1779\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 387px;\"\u003e\n \u003cp\u003e\u003cem\u003eSpiranthes spiralis\u003c/em\u003e (L.) Chevall., 1827\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 115px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003cem\u003eComparison of the flora of cemeteries with the flora of other urban habitats\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe number of indicator species of cemeteries was 288, of which 26 are species of conservation interest. The number of indicator species in other urban habitats was 73, none of which are species of conservation interest. Indicator species lists are in Online Resource 4.\u003c/p\u003e\n\u003cp\u003eThe proportion of native and naturalized species between the two groups of indicator species did not differ significantly (\u0026chi;\u0026sup2; test, p-value = 0.45); around 80% of the indicator species in either context were native (Fig. 6).\u003c/p\u003e\n\u003cp\u003eBoth in cemeteries and in other urban habitats, a significant proportion of indicator species are linked to anthropogenic vegetation. Nevertheless, the proportion of indicator species between the seven habitat types is significantly different (Fisher test, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0,001) (Fig. 7). All species of heathland and dry neutrophilous or calcareous grasslands, as well as the vast majority (3 \u003cem\u003evs.\u003c/em\u003e 67) of species of mesophilous grasslands, are found only among the indicator species of cemeteries. Conversely, species associated with wet herbaceous vegetation and forest vegetation are overepresented among the indicator species of other urban areas.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eThe effect of the urban-rural gradient on the flora of cemeteries\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe number of species whose distribution is significantly linked to cemeteries in an urban context is 148, of which three are of conservation interest. The number linked to cemeteries in a rural context is 130, of which 14 are of conservation interest. The detailed lists can be consulted in Online Resource 5.\u003c/p\u003e\n\u003cp\u003eThe vast majority of species whose distribution is linked to rural cemeteries are native (Fig. 8): only three are naturalized non-natives, whereas this proportion is 38% for species linked to cemeteries in an urban context (\u0026chi;\u0026sup2; test, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001).\u003c/p\u003e\n\u003cp\u003eThere are significant differences in the habitat preference of species associated with cemeteries in urban and rural environments (Fisher\u0026apos;s exact test, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.001) (Fig. 9). Species associated with heathland, and dry neutrophilous or calcareous grasslands and meadows are over-represented in species associated with rural cemeteries, whereas those associated with forests and margins and anthropogenic environments are much more common in species associated with urban cemeteries.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eMany studies on biodiversity in cemeteries have focused on cemeteries that are remarkable for their size, architecture or history (Buchholz et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Buldrini et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Kowarik et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Otves, Arsene et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Šilc \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Yılmaz et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The present study focuses on a synthesis of the floristic biodiversity of cemeteries across an entire region with inventories available for half of the cemeteries in this region, \u003cem\u003ei.e.\u003c/em\u003e 817 cemeteries. Contrary to many studies in which inventories are carried out on a single visit (L\u0026ouml;ki et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019b\u003c/span\u003e; Konic et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), the present dataset covers the entire vegetation period (April to October), which makes it possible to draw up a fairly exhaustive portrait of the flora that develops in the cemeteries of \u0026Icirc;le-de-France, whatever its phenology. In this way, vernal species such as those associated with dry grasslands on acidic sands, and naturalised species, many of which tend to appear late in the season, are not overlooked. Our results may be nuanced by the fact that our cemetery inventories are not necessarily limited to the strict perimeter of cemeteries, but may include some adjacent areas such as the foot of walls outside the cemetery enclosure, some adjacent streets or the cemetery car park. For this reason, our work concerns the floristic diversity of cemeteries and their adjacent areas. For a regional view of the biodiversity of cemeteries, the inclusion of these adjacent areas is probably negligible since it is unlikely that the species in the adjacent areas are specific to them. From an applied point of view, these adjacent areas are very often public, urbanised and managed by the same services as the cemetery, so it is not inconsistent for them to be included in this work.\u003c/p\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eThe importance of cemeteries for the conservation of plant biodiversity\u003c/h2\u003e\u003cp\u003eThe 731 species observed in the 817 cemeteries surveyed in the \u0026Icirc;le-de-France region represent almost half of the spontaneous flora observed at the same time in the region (around 1,530 species, calculated according to the regional list (Perriat et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2020\u003c/span\u003e)), a proportion comparable to that found to a study in the Rurh Metropolitan Region in Germany (Buch et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Given the small area of cemeteries in the region, this figure is not negligible (Vallet \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The high species richness of cemeteries can probably be explained by the fact that there is an important diversity of micro-habitats in cemeteries: walls, very sunny areas and very shaded areas (for example, at the foot of the perimeter walls), grassy areas and, in some cemeteries, a few trees or shrubs, \u003cem\u003eetc\u003c/em\u003e. In addition, the fact that there is a degree of disturbance (mechanical weeding, trampling, soil disturbance to create new graves, \u003cem\u003eetc.\u003c/em\u003e) can also encourage species diversity (Grime \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Nowińska et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The proportion of non-native species in this cemetery flora pool (16%) is comparable to that of the regional flora (13.5%). This proportion is comparable to that observed in the cemetery of Modena (16.9%) (Buldrini et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and much lower than that observed in Poland (33% on average for 78 cemeteries surveyed) (Nowińska et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, it is always complicated to analyse this type of figure, as it depends on the regional pool of species present, the climate, and a range of other factors like the city size (Pyšek \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Furthermore, in our study, we chose to exclude species that were not spontaneous, including subspontaneous species, \u003cem\u003ei.e.\u003c/em\u003e planted species that only escape very locally. It is not always easy in the field to identify whether the species observed has been planted or not (\u003cem\u003ee.g. Sedum\u003c/em\u003e); it is possible that we have been more restrictive in our study than in those previously cited. However, the place of non-native species also depends on the type of funerary practice (Buldrini et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Nowińska et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Gudžinskas \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). In the \u0026Icirc;le-de-France region, the species planted to decorate graves are often poorly adapted to the local climate, as for example, All Saints' Day Chrysanthemums (\u003cem\u003eChrysanthemum \u0026times;grandiflorum\u003c/em\u003e), begonias as \u003cem\u003eBegonia \u0026times;semperflorens-cultorum\u003c/em\u003e or horticultural cultivars that do not naturalise very well (la \u0026ldquo;pens\u0026eacute;e\u0026rdquo;, \u003cem\u003eViola \u0026times;wittrockiana\u003c/em\u003e). Otherwise, more robust species are often planted in pots (Larramendy \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) or in spaces with thin substrate they also die out during summer droughts. This probably limits the presence of non-native species from horticultural origin in our inventories, even if they are present (\u003cem\u003eSyringa vulgaris\u003c/em\u003e, \u003cem\u003eLaburnum anagyroides\u003c/em\u003e, \u003cem\u003eRobinia pseudoacacia\u003c/em\u003e, \u003cem\u003eetc.\u003c/em\u003e). These species are probably more related to the cemetery's ornamental green spaces than to plants introduced by private individuals to decorate graves. Less than 10% of the non-native species found in cemeteries are considered as invasive aliens in the \u0026Icirc;le-de-France region. The species present are considered to be well-established in the region; for these species, control measures are only recommended in special cases, particularly when these species invade natural habitat with conservation interest, and not in anthropogenic contexts (Wegnez \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn our study, of the 935 species present in cemeteries and/or other urban habitats (streets, wasteland, parks \u003cem\u003eetc.\u003c/em\u003e), 31% are significantly more frequent in cemeteries than other urban habitats. For species of conservation interest, this proportion is 51% (144 species of conservation interest recorded out of the 935 species in this dataset). These figures are quite considerable when compared with the small surface area of cemeteries in relation to other urban environments. Since the proportion of native and non-native species between those that prefer cemeteries and those that prefer other urban habitats does not differ, this may indicate that cemeteries are not a particularly important source for the introduction of non-native species, contrary to the conclusion of Walusiak and Krztoń (\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) in Poland. Nevertheless, of the 288 species found preferentially in cemeteries, 40 are non-native, three of which are considered invasive aliens (\u003cem\u003eAcer negundo\u003c/em\u003e, \u003cem\u003eAilanthus altissima\u003c/em\u003e and \u003cem\u003eParthenocissus inserta\u003c/em\u003e); these species could be introduced or find a favourable habitat in cemeteries although they are not unique locations in that respect. Cemeteries are unique in that they provide habitats for species of dry or mesophilous herbaceous environments (dry grasslands on acid and calcareous soils (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e5\u003c/span\u003e), meadows); conversely, species of woodland or wetland habitats are rarely found in cemeteries. In these urban contexts, these woodland and wetland species are probably associated with urban parks or riverbanks. With the exception of park-like cemeteries, these habitats are not very common in cemeteries. The fact that the flora of cemeteries is linked to dry habitats is not surprising (Buldrini et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Buch et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). What is notable is that cemeteries are home to a pool of species associated with oligotrophic habitats, with both acidic and basic pH levels; these ecological traits are particularly rare among plant species in an urban context (Williams et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). It is therefore important to conserve these species and in particular the semi-natural remnant habitats that may exist in some cemeteries, and are of great interest for the conservation of biodiversity in an anthropogenic environment (L\u0026ouml;ki et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Buch et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eConservation issues differ between town and village cemeteries\u003c/h2\u003e\u003cp\u003eAlthough cemeteries are generally located in urbanised areas, the \u0026Icirc;le-de-France region is a good case of study for assessing the influence of an urban-rural gradient, since there is a highly contrasting gradient in the proportion of built-up areas around cemeteries, from cemeteries with less than 1% of built-up areas to cemeteries with 70% of built-up areas. The former probably correspond to village cemeteries, while those with the most built-up areas are cemeteries in large towns. We find a classic pattern in urban ecology (Hill et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), namely that the city favours the presence of non-native species. It can be seen that the majority of species whose distribution is linked to cemeteries with a highly built-up landscape are non-native species, whereas species linked to low percentages of built-up areas are almost all native. This result does not mean that non-native species are absent from rural cemeteries, but that the non-native species they have are present in cemeteries regardless of the level of urbanisation; in other words, it means that cemeteries in highly urbanised contexts have a pool of non-native species that are not found in cemeteries in villages. This means that it is the landscape context that seems to favour the presence of non-native species in cemeteries, rather than the cemeteries themselves being major points of introduction for non-native species. There are also differences in the habitat preferences of species associated with cemeteries in towns and villages. For example, the species assemblages of dry or mesophilous herbaceous environments (acid and limestone grasslands, meadows, heathlands), which have already been mentioned because some of their species are of conservation interest, are more closely associated with rural cemeteries. There are two possible reasons for this (Trzaskowska and Karczmarz \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2013\u003c/span\u003e): urban cemeteries were probably established in areas yet influenced by human activities at the urban fringe with few remnant natural habitats. Morevore, the organization of urban cemeteries was more often planified by landscape architects with little space to pre-existing spontaneous vegetation whereas in villages pre-existing vegetation has sometimes been conserved (Bertrand \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The species that develop preferentially in urban environments tend to be associated with anthropogenic vegetation or forest vegetation. In the case of the latter, this could be linked to the recent development of so-called park-like cemeteries in towns; with vast areas of urban lawns and wooded parts, their flora is more similar to that of urban parks.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eConclusion about cemeteries management practices\u003c/h2\u003e\u003cp\u003eCemeteries play a particularly important role in preserving plant biodiversity in an anthropogenic context. In terms of conservation, particular attention must be paid to cemeteries, which contain areas of semi-natural remnant habitats; it must be a priority to protect these habitats and their associated species. In relation to this, it might be useful to identify cemeteries with remnant natural habitats to bring these conservation issues to the attention of local authorities. These semi-natural habitats should not be destroyed when the areas reserved for graves are enlarged. Their management should also be adapted: no fertilisation and less frequent mowing, as in many cases these areas are managed as lawns. Annual mowing at the end of the season, if possible with arisings removed, would be sufficient. As these areas are generally located on oligotrophic soils, the development of vegetation is limited. In addition, many of these areas have species that flower quite brightly (\u003cem\u003eCalluna vugaris\u003c/em\u003e and \u003cem\u003eErica cinerea, Scabiosa columbaria\u003c/em\u003e, \u003cem\u003eKnautia arvensis, Armeria arenaria\u003c/em\u003e etc.), which are very interesting from an ornamental point of view. So local managers of cemeteries should be informed of these conservation issues and advised on the type of management to be implemented. Moreover, education of cemetery users to support the acceptance of less intensive management, leading to the presence of spontaneous flora in paths (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e5\u003c/span\u003eF) and the presence of tall (but flowering) vegetation in grassy areas would be valuable.\u003c/p\u003e\u003cp\u003eIn addition to the value of these semi-natural habitats, cemeteries host a range of diverse species that are not often found in other habitats of towns and villages. We encourage the maintenance of the diversity of habitats found in cemeteries for example by keeping old walls and maintaining both sunny and shaded areas. Maintaining the diversity of geological and soil contexts is also a factor of diversity on a regional scale (Buch et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e); when developing cemeteries, local substrates should be favoured over importation of topsoil from outside. However, some recent practices linked to the prohibition of the use of pesticides have raised questions about whether the originality of cemetery floras can be maintained. Park-like cemeteries may host a large number of species, but the species found there are largely the same as those found in parks (species of urban lawns and wooded areas). So, on a city scale, their contribution can be rather redundant with parks already present. New pathway management practices that are developing as a result of the stopping of the use of pesticides raise questions about maintaining the specific characteristics of cemetery floras. The waterproofing of pathways (bitumen, cement) greatly reduces the surface area of favourable habitat; this should be limited to certain main pathways at most, leaving at least the secondary pathways permeable. The transformation of pathways in lawns by sowing of grasses (often dominated by red fescues \u003cem\u003eFestuca\u003c/em\u003e) also raises questions. A number of species characteristic of cemeteries are linked to mineral pathways (sand, alluvium), which provide them conditions of sunshine, dryness and low competition (Buch et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) that they will not find in pathways with dense grass cover. These may be, for example, species associated with dry sandy grasslands or with crops, such as different species of poppies (\u003cem\u003ePapaver rhoeas, P. dubium, P. argemone\u003c/em\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e5\u003c/span\u003e) or field marigold (\u003cem\u003eCalendula arvensis\u003c/em\u003e), or species associated with disturbed environments, such as \u003cem\u003eGalium parisiense\u003c/em\u003e. It therefore seems important to retain mineral pathways and to allow them to be grassed over naturally (without sowing) in order to maintain the originality of the floral communities in cemeteries, whether they are linked to \u0026laquo; ordinary \u0026raquo; or conservation interest flora. We therefore strongly encourage the maintenance of paths that are neither waterproofed nor grassed over, with mechanical weed management.\u003c/p\u003e\u003c/div\u003e"},{"header":"Abbreviations","content":"\u003cdiv class=\"DefinitionList\"\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003efollows the French Taxref version 14 (Gargominy et al. 2020). Taxa were generally processed according to the taxonomic rank of the species. However\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003ein order to avoid bias relating to the uneven recording of critical segregates,apomictic taxa (\u003cem\u003ee.g. Rubus\u003c/em\u003e,\u003cem\u003eTaraxacum\u003c/em\u003e,\u003cem\u003eHieracium\u003c/em\u003e,\u003cem\u003eRosa\u003c/em\u003e,etc.) were aggregated to the genus level or in aggregates\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003ethe same applies to taxa whose taxonomic rank has varied between versions of Taxref (\u003cem\u003ee.g. Arenaria serpyllifolia\u003c/em\u003e and \u003cem\u003eA. leptoclados\u003c/em\u003e which were only subspecies in some Taxref versions). For convenience\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003ewe will still use the terms species throughout this paper.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003ch2\u003eFunding\u003c/h2\u003e\n\u003cp\u003eThis study was initiated thanks to the financial support of \u0026quot;L\u0026apos;Institut Paris R\u0026eacute;gion \u0026ndash; Agence r\u0026eacute;gionale de la biodiversit\u0026eacute; (ARB)\u0026quot;.\u003c/p\u003e\n\u003ch2\u003eCompeting Interests\u003c/h2\u003e\n\u003cp\u003eThe author declare she has any interest in the current study.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eThe datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eJ.V. designed the study, performed the statistical analyses and wrote the manuscript.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eI would like to sincerely thank Oliver Pescott and Herv\u0026eacute; Daniel for their encouragement in publishing this work and for their advice opinion on the first draft of this paper\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdam M, Cormier T and Gobled L (2019). 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Urban For \u0026amp; Urban Green 33:92\u0026ndash;98. https://doi.org/10.1016/j.ufug.2017.10.011\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Footnotes","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003e \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.legifrance.gouv.fr/jorf/id/JORFTEXT000043023130\u003c/span\u003e\u003cspan address=\"https://www.legifrance.gouv.fr/jorf/id/JORFTEXT000043023130\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003e \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.legifrance.gouv.fr/loda/id/JORFTEXT000000865328/\u003c/span\u003e\u003cspan address=\"https://www.legifrance.gouv.fr/loda/id/JORFTEXT000000865328/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003e \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.legifrance.gouv.fr/loda/id/LEGITEXT000006059591/\u003c/span\u003e\u003cspan address=\"https://www.legifrance.gouv.fr/loda/id/LEGITEXT000006059591/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"remnant habitats, nature conservation, species of conservation interest, non-native species, graveyards, urban planning","lastPublishedDoi":"10.21203/rs.3.rs-7448159/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7448159/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e The aim of this work is to assess the importance and the originality of plant vascular biodiversity of cemeteries at a regional scale, compare it with other urbanised habitats and assessing their conservation issues according to an urban-rural gradient. The study was carried out in the \u0026Icirc;le-de-France region (France) with a dataset containing 66,973 records of vascular plants collected in 817 cemeteries between 2000 and 2021. We analysed the preference of species for cemeteries or other anthropogenic habitats using indicator value analysis. To describe species-specific responses to the urban-rural gradient, we employed a niche model analysis coupled with a randomization procedure. We then compared naturalness and habitat preferences between the different species lists obtained by these two methods. Among the 731 spontaneous species were inventoried, 84% of which are native. Of these, 69 can be considered as species of conservation interest; species of anthropogenic communities were well represented but the majority were linked to semi-natural habitats: heathlands, dry grasslands on acidic or calcareous soils and meadows in particular. Species linked to cemeteries are associated with semi-natural habitats, while the species linked to other urban habitats are forest or wetland species. Finally, species specifically linked to cemeteries in an urban context are mainly non-native species. Species of semi-natural herbaceous habitats are much more represented among the species linked to rural cemeteries. When they exist in cemeteries, the conservation of remnant semi-natural habitats should be a priority.\u003c/p\u003e","manuscriptTitle":"What role can cemeteries play in conserving plant biodiversity in different urbanisation contexts?","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-09 13:29:43","doi":"10.21203/rs.3.rs-7448159/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-10-27T16:25:27+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-17T08:09:23+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-01T12:59:32+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"265019676018647698788215458271428229464","date":"2025-09-04T12:34:26+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"163621158496332340858119985165751238103","date":"2025-09-03T18:29:50+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-09-03T03:53:29+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-08-28T02:31:38+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-08-27T22:34:10+00:00","index":"","fulltext":""},{"type":"submitted","content":"Urban Ecosystems","date":"2025-08-24T20:08:42+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"4944ca53-7d95-4658-bd5b-688922c35a09","owner":[],"postedDate":"September 9th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-04-13T16:09:43+00:00","versionOfRecord":{"articleIdentity":"rs-7448159","link":"https://doi.org/10.1007/s11252-026-01939-z","journal":{"identity":"urban-ecosystems","isVorOnly":false,"title":"Urban Ecosystems"},"publishedOn":"2026-04-09 15:57:57","publishedOnDateReadable":"April 9th, 2026"},"versionCreatedAt":"2025-09-09 13:29:43","video":"","vorDoi":"10.1007/s11252-026-01939-z","vorDoiUrl":"https://doi.org/10.1007/s11252-026-01939-z","workflowStages":[]},"version":"v1","identity":"rs-7448159","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7448159","identity":"rs-7448159","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}