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11 January 2022

Two million species catalogued by 500 experts

Mark John Costello

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R. Edward DeWalt

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Thomas M. Orrell

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Olaf Banki

3

Mark John Costello

Nord University, Bodø, Norway.

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R. Edward DeWalt

University of Illinois, Champaign, Illinois, USA.

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Thomas M. Orrell

Smithsonian Institution, Washington DC, USA.

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Olaf Banki

Naturalis Biodiversity Centre, Leiden, the Netherlands.

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More than two million accepted species are now listed in the open-access Catalogue of Life (go.nature.com/3ym3h2g). This achievement addresses a major impediment to the management of biodiversity data by presenting an almost complete index of accepted names and known synonyms.

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Nature 601, 191 (2022)
doi: https://doi.org/10.1038/d41586-022-00010-z

Competing Interests
The authors declare no competing interests.

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11 January 2022

EU Nature Restoration Law needs ambitious and binding targets

Kris Decleer

 ORCID: http://orcid.org/0000-0001-9621-8925

0
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Jordi Cortina-Segarra

 ORCID: http://orcid.org/0000-0002-8231-3793

1
&

Aveliina Helm

 ORCID: http://orcid.org/0000-0003-2338-4564

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Kris Decleer

Research Institute for Nature and Forest, Brussels, Belgium.

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Jordi Cortina-Segarra

University of Alicante, Alicante, Spain.

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Aveliina Helm

University of Tartu, Tartu, Estonia.

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Initiatives by the European Commission to restore the continent’s degraded areas (J. Cortina-Segarra et al. Nature 535, 231; 2016) have proved disappointing. As the United Nations Decade on Ecosystem Restoration gathers momentum, the commission is preparing a law that has legally binding targets. To underscore the urgency, some 1,400 European scientists and 30 expert networks and institutions have signed a declaration by the Society for Ecological Restoration Europe (see go.nature.com/3st6k88).

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Nature 601, 191 (2022)
doi: https://doi.org/10.1038/d41586-022-00011-y

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The broad categories of data included in the repository are summarized in Online-only Table 2 and visualized in Fig. 2. Each category is explained in greater detail below, while full details together with accompanying notes are given in the repository (Database_structure.csv) and in Supplementary File 1. Online-only Table 2 gives an overview of data coverage per category, both across all species and for native species separately. A complete list of data sources is available in Supplementary File 2.Fig. 2Visualization of the attributes presented in the database.Full size imageGeneration of the species listTaxon names listed in the most recent and widely accepted New Flora of the British Isles’ index12 were digitized via the Optical Character Recognition Software ReadirisTM 17 (IRIS). Results from the digitization were transferred into a spreadsheet and obvious recognition errors were fixed. The resulting table contained 5,687 taxa and associated taxonomic authorities. A total of 360 unnamed hybrids were excluded, as well as species noted to have only questionable or unconfirmed records, leaving 5,038 species. Forty-one intergeneric hybrid species, 827 entries relating to (notho)subspecies, (notho)varieties, cultivars and forma were also removed along with 720 named hybrids. Species that were included by Stace12 but which he considered not to be part of the flora (i.e. listed as ‘other species’ and ‘other genera’, e.g. genus Tragus or Coreopsis verticillata) were also excluded. Seven species that were labelled ‘extinct’ in the flora were included as there were indications that the species might be in the process of reintroduction (e.g. Bromus interruptus, Bupleurum falcatum and Schoenoplectus pungens). Extinct native and archaeophyte species without any signs of reintroduction (e.g. Dryopteris remota) are also listed but no additional data are provided and they are not included in calculations of completeness of data (Online-only Table 2). The final number of extant species listed here is therefore 3,209 (comprising 1,468 natives, 1,690 aliens and 51 species with unknown status), plus 18 formally extinct species (natives and archaeophytes not seen in the study region since 1999). Species names and taxonomic authorities were revised according to the 2021 reprint of the New Flora of the British Isles, communicated to us by C.A.S. ahead of publication. Genera with less well-defined species – for example due to apomixis – contain additional information on subgenera, sections, and aggregates, as per Stace12. Since misidentifications are common in these groups, we include a column termed ‘unclear_species_marker’ that allows for these species to be quickly identified and excluded from analyses if appropriate. Such genera are often incompletely listed in our database since most microspecies are not sufficiently well defined.TaxonomyNomenclature of the list was checked by Global Names Resolver in the R package ‘taxize’20,21, using the International Plant Names Index (IPNI)22 as the data source, to remove any digitisation errors. Resolved names were used to determine accepted higher taxonomic hierarchy (family, order) again using taxize, with the National Center for Biotechnology Information (NCBI) database. Species that could not be resolved by the Global Names Resolver or did not yield matches in the NCBI database for their higher taxonomic ranks were manually checked for name matches in the World Checklist of Vascular Plants (WCVP)17. Species within the original species list that were found to be identical to a different spelling in WCVP were retained in the database. In such instances, and when slight spelling differences occurred, the columns ‘taxon_name‘ and ‘taxon_name_WCVP‘ differ. To improve clarity, each species is presented here with its unique identification number according to the WCVP (listed as ‘kew_id’) together with three additional columns (i.e. WCVP.URL, POWO.URL and IPNI.URL) which contain hyperlinks to the freely accessible taxon description websites of the (WCVP)17, Plants of the World Online (POWO)23 and (IPNI)22, respectively. Thus, while the taxon names used in the database correspond to those used by Stace12, changes in the accepted species name since publication can be traced in columns ‘taxonomic_status’ and ‘accepted_kew_id’. The family classification of WCVP follows APG IV24 for angiosperms, Christenhusz et al. (2011)25 for gymnosperms and Christenhusz & Chase (2014)26 for ferns and lycopods.Native statusWe offer three different datasets which describe the status of a species as native or non-native, and its level of establishment in BI. The first is extracted from Stace (2019)12, the second contains the status codes used in PLANTATT10 and the unpublished ALIENATT (pers. comm. author K.J.W.) dataset, and the third is extracted from Alien Plants13. The status from Stace12 and Stace & Crawley13 assigns a species to either native or alien status, with aliens subdivided into archaeophytes and neophytes at different levels of establishment (e.g. denizen, colonist etc., see Online-only Table 1). Status codes from the BSBI can be either AC (alien casual), AN (neophyte), AR (archaeophyte), N (native), NE (native endemic) or NA (native status doubtful).Functional traitsData for five ecologically relevant functional traits (i.e. seed mass, specific leaf area [SLA], leaf area, leaf dry matter content [LDMC] and vegetative height) were downloaded from public data available in the TRY database27 (for specific authors see Supplementary File 1 and Supplementary File 2). Averages were calculated using the available measurements downloaded for each species, excluding rows where the measurement was 0. In addition, the maximum vegetative height for each species is given, where available.Realized niche descriptionRealized niche descriptions based on assessments made on plants living in BI are given in the form of Ellenberg indicator values18, as published in PLANTATT10. Ellenberg indicator values place each species along an environmental gradient (e.g. light or salinity) by assigning a number on an ordinal scale, depending on the species preference for the specific gradient (Online-only Table 2). This information is often used to gain insights into environmental changes based on species occurrences28. For species listed under a previously accepted name in PLANTATT, the information was associated with the accepted synonym in Stace (2019)12. Due to the low coverage of PLANTATT for non-native species included in our list, we additionally include Ellenberg indicator values based on Central European assessments, as made available by Döring29. Each Ellenberg category is listed in a separate column, keeping the information from both data sources separate to avoid confounding of assessments based on two different regions (i.e. Britain and Ireland versus Central Europe).Life strategyTo characterize the life strategy of a species, we used the CSR scheme developed by Grime19, which classifies each species as either a competitor (C), stress tolerator (S), ruderal (R) or a combination of these (e.g. CS, SR). CSR classifications were obtained from the Electronic Comparative Plant Ecology database30. Due to the low coverage of available CSR assessments for species in our database (i.e. data available for just 460 out of 3,209 species) we imputed CSR strategies for a further 981 species using available functional trait data, following the method proposed by Pierce et al.31. The functional leaf traits required for this method – i.e. specific leaf area, leaf area, leaf dry matter content – were obtained from the TRY database27. Pre-existing30 and newly imputed CSR strategies are listed in separate columns.Growth form, succulence and life-formPlant growth form descriptions were obtained from the TRY database27 and filtered for those entries given by specific contributors (Online-only Table 2) to maintain consistent use of growth form categories. Information on whether a species was considered to be a succulent was obtained by screening the entire growth form information obtained from the TRY database for the phrase ‘succulence’ or ‘succulent’.Species life-form categories according to Raunkiaer32 were determined for each species in our dataset with regard to the typical life-form of the species as it grows in BI (pers. comm. M.J.M.C.).Associated biome and originInformation given in the Ecoflora database3 for the biome that each species is associated with was matched to the species names according to Stace12. The recognized biome categories follow Preston & Hill33 and are ‘Arctic montane’, ‘Boreal Montane’, ‘Boreo-Arctic Montane’, ‘Boreo-Temperate’, ‘Mediterranean’, ‘Mediterranean-Atlantic’, ‘Southern Temperate’, ‘Temperate’, ‘Wide Boreal’ and ‘Wide Temperate’.For non-native species, the assumed origin (i.e. the region that plants were most likely to have been introduced to BI from, rather than the full non-BI distribution of a species) was adapted from Stace12 into a brief description of their country or region of origin. In addition, these descriptions were manually allocated to the TDWG level 1 regions listed in the World Geographical Scheme for Recording Plant Distributions (WGSRPD, TDWG)34.Species distributionsDistribution metrics for each species are given as the number of 10-km square hectads in BI with records for the species in question within a specified time window. The data were derived from the BSBI Distribution Database35 and were extracted for each species, dividing the study region into Great Britain (incl. Isle of Man), Ireland and the Channel Islands, as previously partitioned for data available in PLANTATT10. The database was queried using species and hectads for grouping, showing only records ‘matching or within 2 km of county boundary’ and excluding ‘do-not-map-flagged occurrences’. The data were not corrected for sampling bias and should therefore only be used as an indication of trends.Hybrid propensityData on hybridization is provided for 641 species, obtained from the Hybrid flora of the British Isles36 which enumerates every hybrid reported in BI up until 2015 (pers. comm. M.R.B.). Each entry was transcribed manually, and then filtered to exclude (a) hybrids that have been recorded, but not formed in the British Isles, (b) triple hybrids (mainly reported for the genus Salix), (c) doubtful records, (d) hybrids between subspecific ranks, and (e) hybrids where at least one parent is not native (only archaeophytes included). This left 821 hybrid combinations for data aggregation. The metric chosen here is hybrid propensity, which is a per-species metric of how many other species a focal species hybridizes with (sensu Whitney et al., 201037). A scaled hybrid propensity metric is also given which was calculated by weighting the hybrid propensity score by the number of intrageneric combinations for a given genus, to account for the greater opportunities of hybridization in larger genera.DNA barcodesDNA barcode sequences for plant species present in BI are currently available for 1,413 species in our database. The information was derived from a dataset of rbcL, matK and ITS2 sequences compiled for the UK flora generated by the National Botanic Garden of Wales and the Royal Botanic Garden Edinburgh38,39 (pers. comm. L.J. and N.D.V.). The data are given as a hyperlink to the record’s page on the Barcode of Life Data Systems (BOLD40) which includes the DNA barcode sequences as well as scans of the herbarium specimen and information on the sample’s collection. Most species have multiple record pages associated with them, due to the sampling of more than one individual. We include a maximum of three BOLD accessions per species; the full range of individuals sampled can be accessed via the original publications38,39. DNA barcodes are almost exclusively available for native species. Future releases of our database will increase the coverage of the non-native flora significantly. Where species in the BOLD database are attributed to a species name that is considered synonymous with another name in our list, the hyperlink is matched to the latest nomenclature12. 1,421 species have at least one sequence associated with them and 935 species have sequence data for all three sequences (rbcL, matK and ITS2).Genome size and chromosome numbersGenome size data for 2,117 specimens (at least one measurement per species) were obtained from various sources. Measurements for a total of 467 species were newly estimated using plant material of known BI origin, often sourced  from the Millennium Seedbank of the Royal Botanic Gardens, Kew (RBG Kew)41. The measurements were made by flow cytometry using seeds or seedlings and following an established protocol42. Information on the extraction buffers and calibration standard species used are available in the file GS_Kew_BI.csv, along with peak CV values of the measurements as a quality control. Where more than one measurement is reported per species, the measurements were made on plant material from different populations or using different buffers. Previously published data for additional species were obtained from reports on the Czech flora43, the Dutch flora44, and prime values listed in the Plant DNA C-values database45,46. Since significant intraspecific differences in genome size between plant material from different geographical origins have previously been described, predominantly due to cytotype diversity in ploidy level47, genome size measurements from previously published sources were assessed with regard to the origin of the material. The column ‘from_BI_material’ (GS_BI.csv, BI_main.csv) allows users to filter for measurements made on material from BI to exclude a potential bias. The information was obtained from the original publication source of each measurement.Chromosome numbers for 1,410 species (at least one chromosome number per species) determined exclusively from material collected in BI were obtained from an extensive dataset compiled by R.J.G. from various published studies, unpublished theses and personal communications from trusted sources. The counts were made between 1898 and 2017, with a large proportion stemming from efforts to achieve greater coverage of the flora by a team of cytologists based at the University of Leicester and headed by R.J.G. Part of the dataset was previously incorporated into the BSBI’s data catalogue5 but has since undergone revisions to incorporate new information and changes in taxonomy. The dataset contained many measurements at subspecies level which were allocated to the species level taxon in our list. This served to include as much of the often considerable infraspecific variation as possible. Since some species for which chromosome counts have been reported elsewhere are lacking chromosome counts from British or Irish material, they are absent from this dataset. To fill such gaps, we also present chromosome numbers from reports on the Czech flora43, the Dutch flora44, and the Plant DNA C-values database45,46. More

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