The Leipzig Catalogue of Vascular Plants (LCVP)

The Leipzig Catalogue of Vascular Plants (LCVP) – An improved

taxonomic reference list for all known vascular plants

Martin Freiberg

, Marten Winter

Alessandro Gentile

, Alexander Zizka

2,3

, Alexandra Nora

Muellner-Riehl

2,4

, Alexandra Weigelt

1,2

, Christian Wirth

1,2,4,5

1. Systematic Botany and Functional Biodiversity / Botanical Garden, Institute of Biology,

Leipzig University, Johannisallee 21-23, 04103 Leipzig, Germany

2. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, sDiv –

Synthesis Centre, Deutscher Platz 5e, Leipzig 04103, Germany

3. Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands

4. Molecular Evolution and Plant Systematics & Herbarium (LZ), Institute of Biology,

Leipzig University, Johannisallee 21-23, 04103 Leipzig, Germany

5. Max Planck Institute for Biogeochemistry; PO Box 100164, 07701 Jena, Germany.

*corresponding author: Martin Freiberg ([email protected])

Abstract

The lack of comprehensive and standardized taxonomic reference

information is an impediment for robust plant research, e.g. in systematics,

biogeography or macroecology. Here we provide an updated and much

improved reference list of 1,315,479 scientific plant taxa names for all

described vascular plant taxa names globally. The Leipzig Catalogue of

Vascular Plants (LCVP; version 1.0.2) contains 351.176 accepted species (plus

6.160 natural hybrids), within 13.422 genera, 561 families and 84 orders. The

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

LCVP a) contains more information on the taxonomic status of global plant

names than any other similar resource and b) significantly improves the

reliability of our knowledge by e.g. resolving the taxonomic status of

~184.000 taxa names compared to The Plant List, the up to date most

commonly used plant name resource. We used ~4500 publications, existing

relevant databases and available studies on molecular phylogenetics to

construct a robust reference backbone. For easy access and integration into

automated data processing pipelines, we provide an ‘R’-package (lcvplants)

with the LCVP.

Background and summary

Due to substantial progress in the last decade in improving plant taxonomy

with phylogenetic findings, an updated global taxonomic reference list was

urgently required. To date, the most commonly used reference list of vascular

plant taxa names is The Plant List (TPL

), hosted by the Royal Botanic Gardens,

Kew. TPL contains 1,166,081 vascular plants names, including 308,407

accepted names, 304,419 of them angiosperms. ~700,000 names of TPL are

synonyms or other taxonomic ranks (subspecies, varieties, forms), including

227,025 unresolved names. The here presented Leipzig Catalogue of Vascular

Plants (LCVP) updates significantly the global knowledge of plant names not

only compared to TPL (see Table 1) and thus is a major improvement for

global plant research. It is based on existing databases and an additional

4,500 publications (see Supplementary File 3 and 4 for a list of full literature

references ordered by families – as plain text and as bibliography RIS file and

File 5 for a list of abbreviated literature references ordered by individual taxa),

which helped to clarify the status of plant names (i.e. accepted, synonymy,

taxonomic placement; see Methods). In the end, 4059 publications provided

relevant and sufficiently robust additional information, e.g. changes in taxa

names and/or their status. A guiding principle during the compilation of the

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

LCVP was to avoid polyphyletic genera, which are frequent in TPL, either by

splitting genera (e.g. separating Goeppertia from Calathea) or fusing them

(e.g. Stapelia and Duvalia in Ceropegia). However, we did not recombine any

species name in the LCVP and in cases of unclear phylogenetic position of

genera, we used the conservative (i.e. existing) name.

Taxonomists, ecologists and conservation biologists often work with many

species (names) and cannot keep pace with the rapid progress in (plant)

systematics, boosted by molecular phylogenetic methods

. These researchers

often rely on taxonomic reference lists as tools to translate taxa names to

accepted species names via accepted synonyms.

Comprehensive taxa lists, such as the LCVP, are essential to standardize

taxonomic names in databases compiled from various sources, relying on a

robust ‘translation’ of species names into one scheme. The TRY database of

functional plant traits (TRY

; www.try-db.org) is one of the most prominent

examples containing trait information for about 150,000 vascular plant

species. Other global databases using plant name reference lists focus on

plant co-occurrence patterns, such as sPlot containing about 1,1 million

vegetation surveys (

~55,000 species), or use any plant species occurrence

information, such as the Global Biodiversity Information Facility (GBIF

~315,000 vascular plant species; www.gbif.org), the Botanical Information

and Ecology Network (BIEN

: ~348,000). The Global Inventory of Floras and

Traits (GIFT

: ~268,000; http://gift.uni-goettingen.de/home) or the inventory

of the Global Naturalized Alien Flora (GloNAF

~14,000; glonaf.org) focus on

plant distribution information from regional floras or floristic inventories.

Generally, such databases were compiled from heterogeneous data sources

varying in time of publication and place of origin. The underlying sources may

be primary or secondary literature-based using work of scientists with

excellent to no plant taxonomic background, thus combining data with

various degrees of complexities and uncertainties. The merging of these

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

databases works via species identities and thus depends on the use of

accepted species names. These databases typically tap phylogenetic

information contained in taxonomic references lists via available tools

supporting automated matching and error checking (i.e. taxon scrubbing).

There is a variety of R packages (e.g. taxonstand

; taxize

; RBIEN

) or online

tools (e.g. Global Name Resolver http://resolver.globalnames.org/ or the

Taxonomic Name Resolution Service

http://tnrs.iplantcollaborative.org/TNRSapp.html) supporting researchers to

check their taxonomic information (see

for a review on some of those tools).

Most of these tools rely on TPL as part of their reference lists, which, however,

has not been updated for a decade and originated in a time when

phylogenetic information on many genera did not exist.

Global taxonomic name databases are useful in their own right, and jointly

create synergies that have transformed ecology into a synthetic and global

science, and can help identifying knowledge gaps

. For example, functional

biogeography combines information on community composition, plant

species distribution and functional traits of the component species to make

inferences on determinants of global trait distribution

. While there is high

potential for exciting research using most up-to-date taxonomic information,

it can be only as good as the input data and the ability of the user to

understand the advantage and shortcoming of the data coming from those

resources. For example, missing taxonomic background often leads to

neglecting the importance of citing authors of names and inevitably leads to

inconsistencies when data from different sources are matched. LCVP shows

that when matching plant taxa names without author names, results could

have up to 10% mismatches (i.e. ~10% of all LCVP plant taxa names are the

same but ultimately refer to different accepted plant taxa).

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

Methods

The creation of the LCVP involved three major steps. (1) We did a thorough

search of available and relevant plant taxonomic databases (Supplementary

File 1) to collate a raw data table of existing plant names (see Step 1:

Producing the raw data table). This table included many contradictory

opinions in taxonomic placement of species. (2) Based on additional

information in ~4500 publications and the reliability, timeliness and quality

of relevant scientific evidences in this literature we decided for each name,

whether that name is in LCVP accepted, synonymous or unresolved (see for

more details Step 2: Decision making). Additionally, we harmonized and

corrected taxa names orthographically. (3) We implemented the LCVP in an

R package (LCVP) which is accessible under a MIT license from GitHub

(https://github.com/idiv-biodiversity/LCVP) and will ensure a coherent

versioning of the list and future updates. Furthermore, we provide a utility

function to use LCVP for taxonomic name resolution (lcvplants), which is also

available under the same license from GitHub (https://github.com/idiv-

biodiversity/lcvplants).

Step 1: Producing the raw data table

TPL provided the core of the basic raw data table for published vascular plant

names, primarily supplemented by the International Plant Names Index

(IPNI

, https://www.ipni.org/). IPNI provides a list of published names and

their source, but does not provide any information on accepted or

synonymous names. We used additional major and minor databases (

17-55

)

which we have chosen based on their availability, on our expert judgement

on comprehensiveness, and whether they contained information if taxa

names are accepted or not (see Supplementary File 1 for a table of used

databases). All additional names and potential synonyms found in those

databases were incorporated in the raw data table.

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

Step 2: Decision making

The extensive raw data table of more than two million entries of plant taxa

names contained a high number of orthographic errors, inconsistencies and

contradictory opinions concerning the status of the names. A rough guideline

for the acceptance of names was a subjective assignment of quality and

reliability to the source. Generally, changes were only applied when the

authors of the respective publications were clearly suggesting those changes.

We ascribed a higher reliability rank (e.g. for conflicting information) usually

to the most recent publications. Additionally, when conflicting information

appeared we usually used information from publications with a) a more

thorough literature section and b) a more comprehensive synonymy history

than to those without. A complete synonymy history should include and

properly cite not only the latest accepted taxon, but also the depending

taxonomic history of all names connected to this taxon (e.g. if it is a

recombined taxon) with all homonymic (i.e. species epitheton is the same)

and heteronymic (i.e. genus name is the same) synonyms. Since phylogenies

based on morphological data alone are prone to homoplasy, only

phylogenetic studies that made taxonomical decisions also based on

molecular data were taken into account. We did not create new species name

combinations. In case of conflicting evidence on the phylogenetic placement

or species name, due to e.g. different methods to build phylogenetic trees,

species names were marked “comb.ined.” following the basionym author.

The following examples illustrate how we treated name changes: The genus

Dracaena and Sansevieria are closely related

, where Sansevieria seems to

be clearly nested within Dracaena, but the differences between both genera

are fluently. Lu et al.

separated the Hawaiian species of Dracaena in a new

genus Chrysodracon, but did not recombine Sansevieria with Dracaena yet.

The presented argumentation and data in

were thorough and

comprehensive and thus we accepted the authors arguments, kept

Sansevieria and Dracaena as distinct genera and separated the Hawaiian

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

species of Dracaena in the new genus Chrysodracon. In another case

Borchsenius et al.

clearly showed that Calathea in the traditional description

was polyphyletic. In order to keep Ischnosiphon and Monotagma as distinct

genera, being the sister clade to a smaller Calathea clade including the type

species, the larger clade of Calathea was put into the then resurrected genus

Goeppertia. The argumentation and presentation in

was robustly based on

a molecular phylogeny producing well supported clades. As consequence we

accepted the recombination of the much larger clade as suggested in

Supplementary Files 3 and 4 provide a complete list of all ~4,500 literature

references ordered by plant families that we used to decide upon species

names to create LCVP. We focused on literature published from 1994

onwards, when molecular phylogenies became widespread (

58,59

Supplementary File 5 provides a list directly matching >104.000 individual

taxa and literature, used to inform the applied name changes for the

respective taxa.

We also applied changes to the spelling of species names. Generally, we

recommend to check the species names prior to automated list treatments,

following the guidelines given in

and the rules of the current version of the

International Code of Nomenclature for algae, fungi, and plants (Shenzhen

Code

). We followed the Shenzen Code using standardized orthography of

epitheta across genera and families, e.g. warscewiczii (neither warscewitzii nor

warszewiczii). Only upper cases from ‘A’ to ‘Z’, lower cases from ‘a’ to ‘z’ and

the hyphen ‘-‘ should be used in the scientific names, special characters are

not valid and to be avoided (Isoëtes- > Isoetes, Köberlinia -> Koeberlinia).

Authors were given in their short form as provided by IPNI. For further

standardization and easier use in automated workflows, we omitted spaces

within author names (C. F. W. Meissn., C.F. W. Meissn., C. F.W. Meissn. C. F.

W.Meissn. -> C.F.W.Meissn.; Balf. f.-> Balf.f.). We linked names published by

two authors with the ‘&’ sign (e.g. Primula minor Balf.f. & Kingdon-Ward).

Names published by three and more authors were restricted to the first

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

authors followed by ‘& al.’ (e.g. Limonium irtaense P.P.Ferrer, A.Navarro,

P.Pérez, R.Roselló, Rosselló, M.Rosato & E.Laguna -> Limonium irtaense

P.P.Ferrer & al.). This refers to the recommendation of the Shenzhen Code,

Art. 46 c. We tried to include only natural hybrids (i.e. no cultivars; based on

expert judgement of LCVP authors) in the LCVP. Since hybrids were not the

focus of the LCVP, we only marked them with ‘_x’, either following the genus

name or the epitheton to recognize them as such, but we did not give any

parent taxa information.

In most cases, we adopted the names used by the taxonomic expert (i.e.

reference author who is usually a person with a publication record within a

certain taxonomic group). However, there are many taxa belonging to genera

or species which have not been phylogenetically analyzed yet. For those, we

adapted the most frequently used taxon name from the recent literature (see

Supplementary Files 3 -5). Despite a major effort, there are still names, which

we could not resolve.

Step 3: Implementation in R

Besides providing LCVP as downloadable tables, we also provide the LCVP as

an R package (LCVP) for easy integration with analyses pipelines. We also

provide a tool for fuzzy-matching-based taxonomic name resolution directly

linked with LCVP (lcvplants). This fuzzy-matching algorithm is applied at

species, infra-species and authority level of a plant taxon; it uses the

‘max.distance’ argument from the agrep() function’ to assess the comparison

between the searched plant name and the closest plant name from the LCVP

list (in terms of number of the same character and their order). The taxonomic

names resolution is implemented in a user-friendly way, and can be done

with few lines of code:

```

# install LCVP and lcvplants from GitHub

install.packages(“devtools”)

library(devtools)

devtools::install_github("idiv-biodiversity/LCVP")

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

devtools::install_github("idiv-biodiversity/lcvplants")

# load the package

library(lcvplants)

# run analyses

LCVP("Hibiscus vitifolius")

```

We provide a description of the fuzzy matching algorithm and its

implementation in Supplementary File 2 and as detailed tutorial on how to

use lcvplants online (https://idiv-biodiversity.github.io/lcvplants/).

Data Records

LCVP contains 1,315,479 vascular plant taxa names. There are 351,176

accepted species names in the LCVP. The accepted species in LCVP belong to

13,422 genera, 561 families, and 84 orders, respectively. LCVP significantly

reduced the number of unresolved plant taxa names by ~184,000 to ~ 60,000

(5%) taxa (see Table 1). The LCVP version 1.0.2 is available in both Microsoft

Excel and text formats in the iDiv data portal

(https://idata.idiv.de/ddm/Data/ShowData/1806). A developmental version

of the LCVP and the lcvplants package are publicly available via GitHub

(https://github.com/idiv-biodiversity/lcvplants). LCVP version 1.0.2 has a DOI

(10.25829/idiv.1806-40-3009 via the iDiv data portal). We will constantly

curate the LCVP and plan to release a new version once every second to third

year.

Technical Validation

We tested whether all synonyms lead to an accepted name or another

synonym. One major issue with TPL is the high amount of unresolved taxa. A

link to another name sometimes is another synonym leading to unresolved

loops. LCVP only links to accepted names, not to the taxonomic predecessor.

If taxon A is synonym to taxon B and it turned out, that taxon B is synonym

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

to taxon C, the accepted name given for taxon A is taxon C, not B. We treated

invalid names as synonyms and assigned them to their appropriate accepted

name.

Most of the still unresolved species names in LCVP were originally published

in the 19th century. There is a high probability that the majority of them are

synonyms, e.g. because of historic transfer errors from one publication to the

other. An extraordinarily high amount of unresolved taxa can be found in

Asteraceae (Hieracium 5,800 names out of a total of 19,300 names are

unresolved, Senecio 685 out of 6,680, Cirsium 357 out of 2,170), Rosaceae

(Rubus 4,040 out of 10,200, Rosa 2,300 out of 5,970, Prunus 512 out of 2,070,

Potentilla 728 out of 3,950, Crataegus 730 out of 2,700, Pyrus 379 out of

1,200), Salicaceae (Salix 619 out of 3,800), Araceae (Anthurium 585 out of

2,260), and Geraniaceae (Pelargonium 963 out of 1,840).

Comparison to TPL

Due to the improved name resolution and increased name information in

general in LCVP compared to TPL, any work flow including taxonomic

harmonization of plant names, will very likely yield more robust and reliable

results for e.g. species richness pattern, matches between different data

sources. For an easier comparison and guidance for users on the differences

between LCVP and TPL, LCVP includes information whether taxa name entries

are identical, differ in the cross-reference to a synonym, differ only

orthographically either by the name or the author, or whether a name is new

in the LCVP and not present in TPL. This unique information makes it possible

for the users of TPL to update their names according to the LCVP, because all

differences are clearly stated in the column ‘status’ of the LCVP.

TPL

LCVP

Plant taxa names

1,166,038

1,315,479

Accepted vascular plant species names

308,397

351,176

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

Synonyms

597,543

679,571

Accepted infraspecific taxa names

20,719

48,257

Accepted Genera

12,660

13,422

Accepted Families

473

561

All Genera

22,830

32,986

Unresolved taxa names

244,017

59,837

TPL - LCVP comparison

Number of records

Identical taxa names

825,173

Additional taxa names in LCVP not being

in TPL

149,474

Families in LCVP not being in TPL

Families in TPL not being in LCVP

Genera in LCVP not being in TPL

2,716

Genera in TPL not being in LCVP

1,954

Resolved in LCVP - unresolved in TPL

182,771

Accepted in LCVP - Synonym in TPL

59,837

Synonym in LCVP - Accepted in TPL

178,673

Different authors in LCVP

31,117

Different synonym in LCVP

65,631

Different orthography in LCVP

6,290

Table 1: Summary of information content in the LCVP and taxa name

differences between LCVP and TPL.

Kew Gardens´ research effort to standardize plant names recently focuses on

their new flagship program, Plants of the World Online (POWO,

http://www.plantsoftheworldonline.org/), which includes a not yet published

new taxonomic reference backbone (Alan Paton from Kew Gardens, pers.

comm.). Given that this is becoming the successor of TPL (see

http://www.plantsoftheworldonline.org/about) we also compared the

available POWO list with LCVP (POWO access date: November 2018; provided

by Kew). With ~335,000 accepted species names and ~458,000 taxa of

vascular plants marked as synonyms in POWO, LCVP contains also

significantly more species name information than POWO (this comparison

includes only vascular plants and excludes infra-specific taxa since LCVP

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

covers only vascular plants and POWO does not include taxa below species

level).

TPL and POWO cover all plants, LCVP only vascular plants. With the current

information we have, LCVP contains more information about vascular plant

taxa names (e.g. more resolved taxa, more accepted species, more synonyms)

than TPL and POWO. A user is more likely to resolve a given vascular plant

taxa name with LCVP than with the given versions of TPL and POWO. LCVP

covers also infraspecific taxa names which are not covered in POWO yet. The

information in LCVP to which genus a species belongs and/or thus which

accepted name should be used, is based on taxonomic, but also on most

recent phylogenetic (i.e. mainly genetic) information. TPL was not updated

for many years, and is mainly based on taxonomic information (i.e. not

molecular phylogenies). With respect to usability of LCVP, we do see

advantages compared to POWO, which to our knowledge does not offer an

R package nor any other functionality of (half)automatic name checking or

any fuzzy name matching functions.

Code Availability

The LCVP generally consists of (1) the LCVP itself available as R data package

(version 1.0.2 as of April 2020) and as tab-delimited textfile file and (2) the R-

package lcvplants. The static LCVP file and the lcvplants tools are publicly

available either via the iDiv data repository:

https://idata.idiv.de/ddm/Data/ShowData/1806 or as developemental

versions via GitHub (https://github.com/idiv-biodiversity/LCVP,

https://github.com/idiv-biodiversity/lcvplants). We plan to closely

collaborate with plant synonymy services and tools like e.g. BIEN, GNR, R

packages taxonstand and taxize, to include LCVP as reference option.

Requests for integrating LCVP can be made via the projects GitHub

(https://github.com/idiv-biodiversity/LCVP/issues)

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

Acknowledgements

We thank thousands of experts working sometimes their whole life on plant

taxonomy and systematics, creating invaluable information and knowledge

the LCVP is also based on. Without this knowledge the LCVP would not exist.

We also thank Anke Stein, Patrick Weigelt, Aldo Compagnoni, Jitendra

Gaikwad, Jens Kattge & Ingolf Kühn for helpful comments on the draft and R

package functionalities. MW, AZ & AG thank DFG for funding (via iDiv, FZT

118). ANMR thanks BMBF for funding (Grant no. 16GW0120K).

Author contributions

MF compiled the LCVP. AG, MW & AZ designed the R packages. AG & AZ

implemented the R functions based on discussions with MW & MF. MW &

MF compiled the drafts of the data paper. All authors contributed to the

writing of the manuscript.

Competing interests

The authors declare no competing interests.

References

1 The Plant List. http://www.theplantlist.org/ (2013)

2 Rouhan, G. & Gaudeul, M. Plant Taxonomy: A Historical Perspective,

Current Challenges, and Perspectives. Methods Mol Biol 1115, 1-37,

doi:10.1007/978-1-62703-767-9_1 (2014).

3 Kattge, J. et al. TRY - a global database of plant traits. Global Change

Biology 17, 2905-2935, doi:10.1111/j.1365-2486.2011.02451.x (2011).

4 Bruelheide, H. et al. sPlot - A new tool for global vegetation analyses. J

Veg Sci 30, 161-186, doi:10.1111/jvs.12710 (2019).

5 GBIF.org. https://www.gbif.org (2019)

6 Enquist, B., Condit, R., Peet, R., Schildhauer, M. & Thiers, B.

Cyberinfrastructure for an integrated botanical information network to

investigate the ecological impacts of global climate change on plant

biodiversity. PeerJ Preprints 4 (2016).

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

7 Weigelt, P., König, C. & Kreft, H. GIFT – A Global Inventory of Floras and

Traits for macroecology and biogeography. Journal of Biogeography 0,

doi:10.1111/jbi.13623.

8 van Kleunen, M. et al. The Global Naturalized Alien Flora (GloNAF)

database. Ecology 100, e02542, doi:10.1002/ecy.2542 (2019).

9 Cayuela, L., Granzow-de la Cerda, I., Albuquerque, F. S. & Golicher, D. J.

TAXONSTAND: An R package for species names standardisation in

vegetation databases. Methods Ecol Evol 3, 1078-1083,

doi:10.1111/j.2041-210X.2012.00232.x (2012).

10 Chamberlain, S. & Szöcs, E. taxize: taxonomic search and retrieval in R.

F1000Research 2, 191, doi:10.12688/f1000research.2-191.v2 (2013).

11 Maitner, B. S. et al. The BIEN R package: A tool to access the Botanical

Information and Ecology Network (BIEN) database. Methods Ecol Evol 9,

373-379, doi:10.1111/2041-210x.12861 (2018).

12 Boyle, B. et al. The taxonomic name resolution service: an online tool

for automated standardization of plant names. Bmc Bioinformatics 14,

16, doi:10.1186/1471-2105-14-16 (2013).

13 Wagner, V. A review of software tools for spell-checking taxon names

in vegetation databases. J Veg Sci 27, 1323-1327, doi:10.1111/jvs.12432

(2016).

14 Cornwell, W. K., Pearse, W. D., Dalrymple, R. L. & Zanne, A. E. What we

(don't) know about global plant diversity. Ecography 0,

doi:10.1111/ecog.04481 (2019).

15 Bruelheide, H. et al. Global trait-environment relationships of plant

communities. Nat Ecol Evol 2, 1906-1917, doi:10.1038/s41559-018-

0699-8 (2018).

16 The Royal Botanic Gardens, Kew, Harvard University Herbaria &

Libraries and Australian National Botanic Gardens. International Plant

Names Index (IPNI) http://www.ipni.org (2010)

17 Clayton, W. D., Vorontsova, M. S., Harman, K. T. & Williamson, H.

GrassBase - The Online World Grass Flora

http://www.kew.org/data/grasses-db.html (2006)

18 The Integrated Taxonomic Information System (ITIS)

https://www.itis.gov/ (2010)

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

19 Pelser, P. B., Barcelona, J. F. & Nickrent, D. L. Co’s Digital Flora of the

Phillippines http://www.philippineplants.org/ (2011)

20 Botanical Garden of Geneva. The African Plant Database

http://www.ville-ge.ch/musinfo/bd/cjb/africa/recherche.php? (2012)

21 Kiefer, M. et al. BrassiBase: Introduction to a Novel Knowledge Database

on Brassicaceae Evolution. Plant and Cell Physiology 55, e3-e3,

doi:10.1093/pcp/pct158 (2013).

22 Dipterocarpaceae Data Base http://rbg-web2.rbge.org.uk/diptero/

(2015)

23 Gentian Research Network

http://gentian.rutgers.edu/classNEW123.htm (2015)

24 Gesneriaceae Research

http://botany.si.edu/gesneriaceae/checklist/result.cfm (2015)

25 SysTax – ein Datenbanksystem für Systematik und Taxonomie

http://www.systax.org/ (2015)

26 M., C., Stanberg, L. & Stevenson, D. The World List of Cycads, online

edition (version Jun 2015). in Species 2000 & ITIS Catalogue of Life, 2018

Annual Checklist (eds Y. Roskov et al.) (Species 2000: Naturalis, 2015)

www.catalogueoflife.org/annual-checklist/2018.

27 Flann, C (ed.). Global Compositae, Checklist Accessed 2016-2019

https://compositae.landcareresearch.co.nz/ (2016)

28 Daniel L. Nickrent (ed.). Parasitic Plant Classification

https://parasiticplants.siu.edu/ListParasites.htm (2016)

29 South African National Biodiversity Institute. Botanical Database of

Southern Africa (BODATSA) http://posa.sanbi.org/sanbi (2016)

30 Vattakaven, T. et al. India Biodiversity Portal: An integrated, interactive

and participatory biodiversity informatics platform. Biodiversity Data

Journal 4, e10279 (2016).

31 International Carnivorous Plant Society. Carnivorous Plant Database

http://www.omnisterra.com/bot/cp_home.cgi (2017)

32 The Flora Malesiana https://floramalesiana.org/new/ (2018)

33 Aedo, C. RJB Geranium: Geranium Taxonomic Information System

(version Sep 2017) in Species 2000 & ITIS Catalogue of Life, 2018 Annual

Checklist (eds Y. Roskov et al.) (Species 2000: Naturalis, 2018)

www.catalogueoflife.org/annual-checklist/2018.

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

34 Culham, A. & Yesson, C. Droseraceae Database (version 0.1, Dec 2008)

in Species 2000 & ITIS Catalogue of Life, 2018 Annual Checklist (eds Y.

Roskov et al.) (Species 2000: Naturalis, 2018)

www.catalogueoflife.org/annual-checklist/2018.

35 Farjon, A., Gardner, M. & Thomas, P. Conifer Database (version Jan

2014) in Species 2000 & ITIS Catalogue of Life, 2018 Annual Checklist

(eds Y. Roskov et al.) (Species 2000: Naturalis, 2018)

www.catalogueoflife.org/annual-checklist/2018.

36 Govaerts R. (ed). WCSP: World Checklist of Selected Plant Families

(version Aug 2017) in Species 2000 & ITIS Catalogue of Life, 2018 Annual

Checklist (eds Y. Roskov et al.) (Species 2000, 2018)

www.catalogueoflife.org/annual-checklist/2018.

37 Hassler, M. World Ferns: Checklist of Ferns and Lycophytes of the World

(version Apr 2018) in Species 2000 & ITIS Catalogue of Life, 2018 Annual

Checklist (eds Y. Roskov et al.) (Species 2000: Naturalis, 2018)

www.catalogueoflife.org/annual-checklist/2018.

38 Hassler, M. World Plants: Synonymic Checklists of the Vascular Plants

of the World (version Apr 2018) in Species 2000 & ITIS Catalogue of Life,

2018 Annual Checklist (eds Y. Roskov et al.) (Species 2000: Naturalis,

2018) www.catalogueoflife.org/annual-checklist/2018.

39 Koch, M. A., German, D. A., Kiefer, M. & Franzke, A. Database

Taxonomics as Key to Modern Plant Biology. Trends in Plant Science 23,

4-6, doi:10.1016/j.tplants.2017.10.005 (2018).

40 Maslin, B. WWW: World Wide Wattle (version 2, Jan 2018) in Species

2000 & ITIS Catalogue of Life, 2018 Annual Checklist (eds Y. Roskov et

al.) (Species 2000: Naturalis, 2018) www.catalogueoflife.org/annual-

checklist/2018.

41 Rainer H. & Chatrou L.W. (eds). AnnonBase: Annonaceae GSD (version

Jan 2014) in Species 2000 & ITIS Catalogue of Life, 2018 Annual Checklist

(eds Y. Roskov et al.) (Species 2000: Naturalis, 2018)

www.catalogueoflife.org/annual-checklist/2018.

42 Roskov, Y., Zarucchi, J., Novoselova, M. & Bisby F. (eds). ILDIS World

Database of Legumes (version 12, May 2014) in Species 2000 & ITIS

Catalogue of Life, 2018 Annual Checklist (eds Y. Roskov et al.) (Species

2000: Naturalis, 2018) www.catalogueoflife.org/annual-checklist/2018.

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

43 Warwick, S. I., Francis, A. & Al-Shehbaz, I. A. Brassicaceae species

checklist and database (version 2, Oct 2009) in Species 2000 & ITIS

Catalogue of Life, 2018 Annual Checklist (eds Y. Roskov et al.) (Species

2000: Naturalis, 2018) www.catalogueoflife.org/annual-checklist/2018.

44 Royal Botanic Gardens, Kew. Plants of the World Online

http://www.plantsoftheworldonline.org/ (2019)

45 The Chinese Virtual Herbarium (CVH) http://www.cvh.ac.cn/ (2019)

46 Jardim Botânico do Rio de Janeiro. Flora do Brasil 2020

http://floradobrasil.jbrj.gov.br/reflora/listaBrasil/ConsultaPublicaUC/C

onsultaPublicaUC.do#CondicaoTaxonCP (2019)

47 Melastomataceae.Net 2007–2020. A Site with Information on the

Biodiversity of Melastomataceae www.melastomataceae.net (2019)

48 The Smithsonian National Museum of Natural History Botany Collection

https://collections.nmnh.si.edu/search/botany/ (2019)

49 Couvreur, T. L. P. World Annonaceae http://annonaceae.myspecies.info

(2019)

50 Euro+Med. Euro+Med PlantBase

http://ww2.bgbm.org/EuroPlusMed/query.asp (2019)

51 Kilian, N., Hand, R. & von Raab-Straube E. (eds). Cichorieae Systematics

Portal http://cichorieae.e-taxonomy.net/ (2019)

52 Matija Strlic (ed). The Pelargonium Page

http://www.pelargonium.si/index.html (2019)

53 Missouri Botanical Garden. Tropicos.org http://www.tropicos.org (2019)

54 Roskov , Y. et al. Species 2000 & ITIS Catalogue of Life, 2019 Annual

Checklist. (Species 2000: Naturalis, 2019)

www.catalogueoflife.org/annual-checklist/2019.

55 USDA & NRCS. National Plant Data Team. The PLANTS Database

http://plants.usda.gov (2019)

56 Lu, P.-L. & Morden, C. W. Phylogenetic Relationships among

Dracaenoid Genera (Asparagaceae: Nolinoideae) Inferred from

Chloroplast DNA Loci. Systematic Botany 39, 90-104, 115 (2014).

57 Borchsenius, F., Suárez, L. S. S. & Prince, L. M. Molecular Phylogeny and

Redefined Generic Limits of Calathea (Marantaceae). Systematic Botany

37, 620-635, 616 (2012).

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint

58 Soltis, P. S. & Soltis, D. E. Plant Molecular Systematics in Evolutionary

Biology (eds Max K. Hecht, Ross J. Macintyre, & Michael T. Clegg) Ch.

Plant Molecular Systematics - Inferences of Phylogeny and Evolutionary

Processes, 139-194 (Springer US, 1995) Boston, MA.

59 Wolf, P. G., Soltis, P. S. & Soltis, D. E. Phylogenetic-Relationships of

Dennstaedtioid Ferns - Evidence from Rbcl Sequences. Mol Phylogenet

Evol 3, 383-392, doi:DOI 10.1006/mpev.1994.1044 (1994).

60 Stearn, W. T. Botanical Latin. 546 (Timber Press, 2004).

61 Turland, N. J. et al. International Code of Nomenclature for algae, fungi,

and plants (Shenzhen Code) adopted by the Nineteenth International

Botanical Congress Shenzhen, China, July 2017. Vol. 159 (Koeltz

Botanical Books, 2018).

.CC-BY 4.0 International licenseavailable under a

was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made

The copyright holder for this preprint (whichthis version posted May 9, 2020. ; https://doi.org/10.1101/2020.05.08.077149doi: bioRxiv preprint