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Home , Anastatica, Anelsonia, Armoracia, Aubrieta, Boechera, Crucihimalaya

Pl. Syst. Evol. 259: 249–258 (2006) DOI 10.1007/s00606-006-0422-0

Brassicaceae: checklist and database on CD-Rom

S. I. Warwick1, A. Francis1, and I. A. Al-Shehbaz2

1Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Central Experimental Farm, Ottawa, Ontario, Canada 2Missouri Botanical Garden, St. Louis, Missouri, USA

Received October 6, 2005; accepted November 17, 2005 Published online: June 19, 2006 Ó Springer-Verlag 2006

Abstract. A species checklist has been prepared for rank) on the basis of the most recent taxo- the (Cruciferae) family, providing the nomic/floristic treatments for specific regions, first updated list in over 70 years. The family, e.g. Europe (Jalas and Suominen 1994, Jalas et currently, includes 338 genera and 3709 species. al. 1996), North America (Rollins 1993), China The database contains approximately 14,000 taxo- (Zhou et al. 2001), etc. (see literature cited). nomic names (records). Taxon status and synon- Our decisions were also based on the many ymy, taxon name, scientific authority, literature taxonomic changes that have resulted from source and source verification, and the basionym are provided for each record. recent molecular phylogenetic studies. Cur- rently, 338 genera and 3709 species are Key words: Brassicaceae, species checklist, accepted (Table 1). Recent changes include Cruciferae, , database. transfers from to Boechera in North America; from Braya to Xerodraba in South America, from Sisymbrium to Crucihimalaya and Olimarabidopsis in Central Asia, and from Introduction to the database or to Yinshania A species checklist has been prepared for the in China. Within North America, most species Brassicaceae (Cruciferae) family. The last of were found to be better placed comprehensive monographic treatment of the with the earlier-described , family was that of Schulz (1936). The current while one group of auriculate-leaved taxonomic database (Microsoft access format) Lesquerella species was segregated into the contains approximately 14,000 names, genus . As a result, the genus Lesque- compiled from the taxonomic literature start- rella has been placed in synonymy. Other ing with Linnaeus’s in 1753 genera, whose reduction to synonymy is widely to the present and floras from all parts of the accepted, include Dentaria (now Cardamine), world, as well as numerous journal publica- Coronopus (now Lepidium), Guenthera (now tions, including monographs, revisions, and Brassica), Eunomia (now Aethionema), Ero- synopses. The authors have made decisions on phila (now ) and Hutchinsia, Pritzelago the status of all taxa (at least at the specific and Hymenolobus (now Hornungia). 250 S. I. Warwick et al.: Brassicaceae: Species checklist

Table 1. List of genera and accepted number of Table 1. (Continued) species in the Brassicaceae Calepina Adans. 1 Genus No. of Callothlaspi F.K. Mey. 1 species Calymmatium O.E. Schulz 2 Camelina Crantz 11 Acanthocardamum Thell. 1 Camelinopsis A.G. Mill. 1 ´ Achoriphragma Sojak4Capsella Medik. 5 Aethionema R. Br. 56 Cardamine L. 197 Alliaria Heist. ex Fabr. 2 Carinavalva Ising 1 Alyssoides Mill. 2 Carrichtera DC. 1 Alyssopsis Boiss. 2 Catadysia O.E. Schulz 1 Alyssum L. 195 Catenulina Soja´ k1 Ammosperma Hook. f. 2 Catolobus (C.A. Mey.) Al-Shehbaz 1 Anastatica L. 1 S. Watson 17 Anchonium DC. 2 Ceratocnemum Coss. & Balansa 1 Andrzeiowskia Rchb. 1 Chalcanthus Boiss. 1 Anelsonia J.F. Macbr. & Payson 1 Chamira Thunb. 1 Aphragmus Andrz. ex DC. 11 Chartoloma Bunge 1 Aplanodes Marais 2 Chaunanthus O.E. Schulz 3 Arabidella (F. Muell.) O.E. Schulz 6 Chilocardamum O.E. Schulz 1 (DC.) Heynh. 11 Chlorocrambe Rydb. 1 Arabis L. 118 Chorispora R. Br. ex DC. 11 Arcyosperma O.E. Schulz 1 Christolea Cambess. ex Jacquem. 2 Armoracia Gaertn., B. Mey. & Scherb. 3 Chrysochamela (Fenzl) Boiss. 3 Aschersoniodoxa Gilg & Muschl. 3 Cithareloma Bunge 3 Asperuginoides Rauschert 1 Clastopus Bunge ex Boiss. 2 Asta Klotzsch & O.E. Schulz ex 2 Clausia Korn.-Trotzky 6 O.E. Schulz Clypeola L. 9 Atelanthera Hook. f. & Thomson 1 Cochlearia L. 19 Athysanus Greene 2 Coincya Porta & Rigo ex Rouy 6 Adans. 12 Coluteocarpus Boiss. 1 Aurinia Desv. 9 Conringia Heist. ex Fabr. 6 Baimashania Al-Shehbaz 2 Cordylocarpus Desf. 1 Ballantinia Hook. f. ex E.A. Shaw 1 Crambe L. 34 Barbarea R. Br. 29 Crambella Maire 1 Berteroa DC. 5 Cremolobus DC. 8 Berteroella O.E. Schulz 1 Crucihimalaya Al-Shehbaz, 9 Biscutella L. 53 O’Kane & R.A. Price Bivonaea DC. 1 Cryptospora Kar. & Kir. 3 Blennodia R. Br. 2 Cuphonotus O.E. Schulz 2 Boechera A. Lo¨ ve & D. Lo¨ ve 65 Cusickiella Rollins 2 Boreava Jaub. & Spach 2 Cymatocarpus O.E. Schulz 3 Bornmuellera Hausskn. 7 Cyphocardamum Hedge 1 Borodinia N. Busch 1 Dactylocardamum Al-Shehbaz 1 Botschantzevia Nabiev 1 Degenia Hayek 1 Brassica L. 39 Delpinophytum Speg. 1 Braya Sternb. & Hoppe 13 Descurainia Webb & Berthel. 48 Brayopsis Gilg & Muschl. 6 Desideria Pamp. 12 Brossardia Boiss. 1 Diceratella Boiss. 11 Bunias L. 3 Dichasianthus Ovcz. & Yunussov 1 Cakile Mill. 6 Dictyophragmus O.E. Schulz 2 S. I. Warwick et al.: Brassicaceae: Species checklist 251

Table 1. (Continued) Table 1. (Continued) Genus No. of Halimolobos Tausch 16 species Harmsiodoxa O.E. Schulz 3 Heldreichia Boiss. 4 Didesmus Desv. 2 Heliophila L. 82 Didymophysa Boiss. 2 Hemicrambe Webb 3 Dielsiocharis O.E. Schulz 2 Hemilophia Franch. 5 Dilophia Thomson 2 Henophyton Coss. & Durieu 2 Dimorphocarpa Rollins 4 Hesperidanthus (B.L. Rob.) Rydb. 5 Diplotaxis DC. 32 Hesperis L. 46 Dipoma Franch. 1 Hollermayera O.E. Schulz 1 Diptychocarpus Trautv. 1 Hormathophylla Cullen & T.R. Dudley 10 Dithyrea Harv. 2 Hornungia Rchb. 3 Dontostemon Andrz. ex C.A. Mey. 11 Horwoodia Turrill 1 Douepea Cambess. ex Jacquem. 2 Hutchinsia R. Br. 1 Draba L. 363 Ianhedgea Al-Shehbaz & O’Kane 1 Drabastrum (F. Muell.) O.E. Schulz 1 Iberis L. 27 Dryopetalon A. Gray 5 Idahoa A. Nelson & J.F. Macbr. 1 Eigia Soja´ k1Iodanthus Torr. & A. Gray ex Steud. 1 Elburzia Hedge 1 Ionopsidium (DC.) Rchb. 5 Enarthrocarpus Labill. 5 Irenepharsus Hewson 3 Englerocharis Muschl. 2 L. 79 Eremobium Boiss. 1 Iskandera N. Busch 2 Eremoblastus Botsch. 1 Ivania O.E. Schulz 1 Eremodraba O.E. Schulz 2 Kernera Medik. 1 Eremophyton Be´ g. 1 Kremeriella Maire 1 Eruca Mill. 4 Lachnocapsa Balf. f. 1 Erucaria Gaertn. 10 Lachnoloma Bunge 1 Erucastrum (DC.) C. Presl 25 Leavenworthia Torr. 8 Erysimum L. 223 Leiospora (C.A. Mey.) F. Dvora´ k6 Euclidium R. Br. 1 Lepidium L. 231 Eudema Humb. & Bonpl. 6 Lepidostemon Hook. f. & Thomson 6 Eunomia DC. 1 Leptaleum DC. 1 Eurycarpus Botsch. 2 Lithodraba Boelcke 1 Eutrema R. Br. 26 Litwinowia Woronow 1 Farsetia Turra 27 Lobularia Desv. 4 Fezia Pit. ex Batt. 1 Lunaria L. 3 Fibigia Medik. 13 Lyrocarpa Hook. & Harv. 3 Foleyola Maire 1 Macropodium R. Br. 2 Fortuynia Shuttlw. ex Boiss. 2 Malcolmia R. Br. 32 Fourraea Greuter & Burdet 1 Mancoa Wedd. 11 Gagria M. Kra´ l1Maresia Pomel 3 Galitzkya V.V. Botschantz. 3 Mathewsia Hook. & Arn. 10 Geococcus J.L. Drumm. ex Harv. 1 Matthiola R. Br. 48 Glastaria Boiss. 1 Megacarpaea DC. 9 Goldbachia DC. 5 Megadenia Maxim. 1 Graellsia Boiss. 8 Menkea Lehm. 6 Grammosperma O.E. Schulz 1 Menonvillea DC. 24 Guillenia Greene 1 Microlepidium F. Muell. 2 Guiraoa Coss. 1 Microstigma Trautv. 3 Gynophora Gilli 1 Microthlaspi F.K. Mey. 4 252 S. I. Warwick et al.: Brassicaceae: Species checklist

Table 1. (Continued) Table 1. (Continued) Genus No. of Phlebolobium O.E. Schulz 1 species Phlegmatospermum O.E. Schulz 4 Nutt. 1 Morettia DC. 3 Physaria (Nutt.) A. Gray 105 Moricandia DC. 8 Physocardamum Hedge 1 Moriera Boiss. 1 Physoptychis Boiss. 2 Morisia J. Gay 1 Physorhynchus Hook. 2 Murbeckiella Rothm. 5 Planodes Greene 1 Muricaria Desv. 1 Polyctenium Greene 2 Myagrum L. 1 Polypsecadium O.E. Schulz 3 Nasturtiopsis Boiss. 1 Pringlea Anderson ex Hook. f. 1 R. Br. 10 Pseuderucaria (Boiss.) O.E. Schulz 2 Neotchihatchewia Rauschert 1 Pseudoarabidopsis Al-Shehbaz, 1 Neotorularia Hedge & J. Le´ onard 11 O’Kane & R.A. Price Nerisyrenia Greene 8 Pseudocamelina (Boiss.) N. Busch 3 Neslia Desv. 1 Pseudoclausia Popov 10 Neuontobotrys O.E. Schulz 5 Pseudofortuynia Hedge 1 N.H. Holmgren 1 Pseudosempervivum (Boiss.) Grossh. 3 Noccaea Moench 77 Pseudoturritis Al-Shehbaz 1 Noccidium F.K. Mey. 1 Pseudovesicaria (Boiss.) Rupr. 1 Notoceras R. Br. 1 Psychine Desf. 1 Notothlaspi Hook. f. 2 Pterygiosperma O.E. Schulz 1 Ochthodium DC. 1 Pugionium Gaertn. 3 Octoceras Bunge 1 Pycnoplinthopsis Jafri 1 Olimarabidopsis Al-Shehbaz, 3 Pycnoplinthus O.E. Schulz 1 O’Kane & R.A. Price Quezeliantha H. Scholz ex Rauschert 1 Onuris Phil. 6 Raffenaldia Godr. 2 Oreoblastus Suslova 1 Raphanoryncha Rollins 1 Oreoloma Botsch. 3 Raphanus L. 3 Oreophyton O.E. Schulz 1 Rapistrum Crantz 2 Ornithocarpa Rose 2 Rhammatophyllum O.E. Schulz 10 Orychophragmus Bunge 2 Rhizobotrya Tausch 1 Otocarpus Durieu 1 Ricotia L. 9 Pachycladon Hook. f. 10 Robeschia Hochst. ex O.E. Schulz 1 Pachymitus O.E. Schulz 1 Rollinsia Al-Shehbaz 1 Pachyneurum Bunge 1 Romanschulzia O.E. Schulz 13 Pachyphragma (DC.) Rchb. 1 Scop. 86 Pachypterygium Bunge 3 Rytidocarpus Coss. 1 Parlatoria Boiss. 2 Sameraria Desv. 9 Parodiodoxa O.E. Schulz 1 Sandbergia Greene 1 Parolinia Webb 5 Sarcodraba Gilg & Muschl. 3 Parrya R. Br. 34 Savignya DC. 1 Paysonia O’Kane & Al-Shehbaz 8 Scambopus O.E. Schulz 1 Pegaeophyton Hayek 7 Schimpera Hochst. & Steud. ex Endl. 1 & Hand.-Mazz. Schivereckia Andrz. ex DC. 2 Peltaria Jacq. 4 Schizopetalon Sims 10 Peltariopsis (Boiss.) N. Busch 2 Schouwia DC. 1 Pennellia Nieuwl. 9 Scoliaxon Payson 1 Petrocallis R. Br. 1 Selenia Nutt. 4 Petroravenia Al-Shehbaz 1 Shangrilaia Al-Shehbaz, 1 Phaeonychium O.E. Schulz 7 J.P. Yue & H. Sun S. I. Warwick et al.: Brassicaceae: Species checklist 253

Table 1. (Continued) Table 1. (Continued) Genus No. of Winklera Regel 3 species Xerodraba Skottsb. 7 Yinshania Ma & Y.Z. Zhao 13 Sibara Greene 9 Zerdana Boiss. 1 Sibaropsis S. Boyd & T.S. Ross 1 Zilla Forssk. 2 Sinapidendron Lowe 4 Zuvanda (Dvora´ k) Askerov 3 Sinapis L. 5 TOTAL No. genera 338 Sisymbrella Spach 2 TOTAL No. species 3709 Sisymbriopsis Botsch. & Tzvelev 5 Sisymbrium L. 95 Skottsbergianthus Boelcke 1 The following information is provided for Smelowskia C.A. Mey. 25 each taxonomic name (record) in the database: Sobolewskia M.Bieb. 4 taxon status and synonymy, taxon name, Solms-laubachia Muschl. 9 scientific authority, literature source and Sphaerocardamum S. Schauer 8 source verification, and the basionym. The Spirorhynchus Kar. & Kir. 1 details of each will be discussed below: Spryginia Popov 7 Taxon status and synonymy – ACC field Stanleya Nutt. 6 (column). Y (currently accepted) or N (not Stenopetalum R. Br. ex DC. 10 accepted), and if N the accepted synonymy is Sterigmostemum M.Bieb. 7 indicated under the ‘‘SYNONYM OF’’ field. Stevenia Adams ex Fisch. 4 A decision has been made on the status of all Straussiella Hausskn. 1 taxa at the specific rank and many, but not all, Streptanthella Rydb. 1 Streptanthus Nutt. 34 of the subspecies and varieties. Subspecies or Streptoloma Bunge 2 varieties of uncertain status or unknown to the Strigosella Boiss. 4 authors are indicated as ‘‘?’’ in the ACC field. Stubendorffia Schrenk. 8 Taxon name. GENUS, SPECIES, SUB- ex Fisch. & Ave´ -Lall. SP_VAR fields. Subularia L. 2 Scientific Authority. BAUTH, AUTH and Succowia Medik. 1 AUTHFULL fields. Author abbreviations and Synstemon Botsch. 2 full names have been standardized in accor- Synthlipsis A. Gray 3 dance with Brummitt and Powell (1992) and in Tauscheria Fisch. ex DC. 1 accordance with the rules of nomenclature. Teesdalia R. Br. 3 According to these rules, if there is clear Tetracme Bunge 10 Thelypodiopsis Rydb. 17 acknowledgment in a given publication that Thelypodium Endl. 19 another author’s manuscript has been used for Thlaspi L. 55 the citation and description of a taxon, the Thysanocarpus Hook. 5 latter author would be the authority. For Trachystoma O.E. Schulz 3 example, Nuttall gave his manuscripts to Transberingia Al-Shehbaz & O’Kane 1 Torrey and A. Gray who acknowledged the Trichotolinum O.E. Schulz 1 use of his descriptions in the introduction to Hook. 4 their Flora of North America and, therefore, Turritis L. 2 Nuttall alone is the author of taxa listed in that Vania F.K. Mey. 1 manuscript. On the other hand, if an author Vella L. 7 has attached another author’s name to a Veselskya Opiz 1 description of a published ‘‘nomen nudum’’ Warea Nutt. 4 or to a previously unpublished name from a Weberbauera Gilg & Muschl. 23 Werdermannia O.E. Schulz 5 herbarium or other source, the cited author will appear as ‘‘ex’’ the author of the publication. 254 S. I. Warwick et al.: Brassicaceae: Species checklist

Literature Source and source verifica- description seen by the authors is indicated by tion. SOURCE and SOURCEAUTH fields. ‘‘y’’, and a second description by ‘‘#’’. The latter field is provided only if different A few of the taxonomic names (with an from the species authority. Periodical citations asterisk ‘‘*’’ after the name) listed in the have been standardized and abbreviated in database are illegitimately published. These accordance with B-P-H., Botanico-Periodi- were included either because they are cited in cum-Huntianum (Bridson 2004, Bridson and the synonymy of major floras and/or have Smith 1991, Lawrence et al. 1968). Transliter- been widely cited over a long period. Taxa ations from Slavic to Roman script, and invalidly published include: 1. unaccompanied numerous name changes and dating of jour- by (or not referring to previously published) nals also follow B-P-H. A number of publica- Latin descriptions or diagnoses on or after 1 tions from parts of the former Soviet Union January 1935, 2. failure to designate a type were written in the original language but specimen starting from 1 January 1958, 3. appeared with an additional Latin title page, failure to include the rank of a new combina- often cited in the literature. These references tion or name on or after 1 January 1953, or 4. are now cited only in the original language of failure to properly cite a basionym and refer- the publication in accordance with current ence on or after 1 January 1953. A name usage. An exception to citing a succession of without description, ‘‘nomen nudum,’’ is often changes in title has been ‘‘Feddes Repertori- cited from an annotated herbarium label or um’’ which we have decided to cite only in this list, and is included in the database only if it more recent simplified form. has been legitimized by another author, in Citations for monographs generally follow which case the latter alone, or rarely both, are Stafleu and Cowan (1976–1988) and Stafleu cited. Such taxa make up the majority of and Mennega (1995). These publications have ‘‘author ex’’ citations. A later homonym of an been particularly useful for checking page earlier published name is illegimate; most of numbers and dates of multi-part monographs, these appear in the literature as ‘‘author non or of preprints or reprints of monographs another author’’ or ‘‘sensu the later author’’ which also appeared in journals. Where pos- after the literature citation. Cases where the sible, the earliest date of publication is cited. If later author amends the original description there is doubt concerning dates, both publica- and applies the amended description to the tions may be cited. synonymy of more than one taxon, are indi- With respect to source verification, a com- cated by ‘‘pro parte.’’ Sometimes the later prehensive search has been made for the homonym applies to a totally different plant, original literature, which has been copied and having been published without knowledge of archived by the authors, and publication the original name, and is treated as ‘‘nomen details verified. Any errors or omissions illeg.’’ A name described only as a synonym of regarding authority names, references, or dates a species, subspecies or variety, is indicated by have been corrected accordingly. Most of the ‘‘pro syn.’’ at the end of the reference. listed species can be found in Index Kewensis, Basionym. BASION field although subspecies and varieties were not The status of many taxa in the Brassicaceae listed there until 1975, and some obscure remains in a state of flux as more information is references were missed. For these, the Gray gathered from all parts of the world, and as an Herbarium Index and the International increasing number of taxa are included in Plant Names Index (IPNI) have also been comparative phylogenetic studies. It is hoped consulted. If an original reference could not be that this database and the accompanying found, it has sometimes been possible to find a database on chromosome counts (Warwick description by the same author in a subsequent and Al-Shehbaz, in this volume) will serve as publication. In the ‘‘AY’’ column, an original reference guides for selecting appropriate taxa S. I. Warwick et al.: Brassicaceae: Species checklist 255 for future taxonomic and phylogenetic studies Bramwell D., Bramwell Z. I. (1974) Wild flowers of within the family. The electronic format allows the Canary Islands, Cruciferae. Stanley Thornes for easy updating of the checklist on a regular Ltd., London, pp. 128–133. basis. We do not claim that our list is complete Bridson G. D. R. (Compiler) (2004) PBH-2, Peri- nor do we feel that it is perfect, and we suspect odicals with botanical content. Hunt Institute that some mistakes may be found. In order to for Botanical Documentation, Carnegie Mellon University, Pittsburgh. keep the list up-to-date, we encourage the Bridson G. D. R., Smith E. R. (eds.) (1991) B-P-H., reader to call our attention to any inaccuracies. Botanico-Periodicum-Huntianum Supplement. Our goal is to continue to serve the Brassica- Hunt Institute for Botanical Documentation, ceae taxonomic community and the others Carnegie Mellon University, Pittsburgh. interested, and we cannot achieve that goal Brummitt R. K., Powell C. E. (eds.) (1992) Authors without feedback. of plant names. Royal Botanic Gardens, Kew London, pp. 1–732. 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