Brassicaceae: Chromosome Number Index and Database on CD-Rom

Pl. Syst. Evol. 259: 237–248 (2006) DOI 10.1007/s00606-006-0421-1

Brassicaceae: Chromosome number index and database on CD-Rom

S. I. Warwick1 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 27, 2005 Published online: June 19, 2006 Springer-Verlag 2006

Abstract. A summary of chromosome numbers has separated and arranged by increasing number, been prepared for the Brassicaceae (Cruciferae) records with the same value were sorted alpha- family. Over 9000 chromosome counts have been betically by country, and lastly by chronological compiled from the literature. Chromosome numbers date of the report. The literature source for each are known from 232 of the 338 (68.6%) genera and count is provided, and includes the source 1558 of the 3709 (42.0%) of the species in the family. (REPORT_BY) if different from the authors of Counts in the excel database have been arranged the publication. The final column for each record according to currently accepted taxonomy, and includes for each count, country of origin and (REPORT_AS) indicates the name of the taxon reference citation. that the count was reported as, if different from the accepted name. The latter synonyms also Key words: Brassicaceae, chromosome number, appear in the alphabetized taxon list (in italics), Cruciferae, chromosome counts, cytology. with the instruction to see the accepted name for information on counts reported under that name. The last column may also indicate if the count has been reassigned by other authors, or Introduction to the database would appear to be erroneous due to misidentif- A summary of chromosome numbers has been ication and/or other errors. All literature cita- prepared for the Brassicaeae (Cruciferae) family. tions are provided in a separate file. Over 9000 chromosome counts have been com- Chromosome numbers are known from piled from literature dating from the first com- 232 of the 338 (68.6%) genera and 1558 of the prehensive cytological studies of Manton (1932) 3709 (42.0%) of the species in the family and Jaretzky (1928, 1932) through to 2005. (summarized in Table 1). A continuous series Chromosome counts in the database (Microsoft of base chromosome numbers from 4 to 13 excel format) have been arranged according to (17) exists, but a surprisingly high percentage the currently accepted taxonomy; see Species (37%) appear to be based on eight. The Checklist for the Brassicaceae in the same issue haploid numbers are often highly variable (Warwick et al.). For each accepted taxon (in even within genera (e.g. Brassica with n =7, bold), haploid (n) and diploid (2n) counts were 8, 9, 10, 11, 16, 17, 18, 19). The lowest number 238 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database

Table 1. Summary for each genus, number of species counted, base chromosome number, and summary of n and 2n chromosome numbers for the genus Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Acanthocardamum Thell. 1 – – – Achoriphragma Soja´k4–– – Aethionema R. Br. 56 16 7, 8 n = 7, 8, 11, 12, 14, 16, 18, 22, 24; 2n = 14, 16, 22, 24, 28, 36, 48, 60 Alliaria Heist. ex Fabr. 2 1 7, 9 n = 7, 18, 21; 2n = 36, 42 Alyssoides Mill. 2 2 8 2n =8,16 Alyssopsis Boiss. 2 1 8 n =8 Alyssum L. 195 103 7, 8 n = 8, 11, 12, 14, 15, 16, 18, 20, 24, 32; 2n = 14, 16, 24, 28, 30, 32, 46, 48, 64, 88, 92 Ammosperma Hook. f. 2 – – – Anastatica L. 1 1 11 2n =22 Anchonium DC. 2 1 7, 8 n =7;2n = 14, 16, 21 Andrzeiowskia Rchb. 1 – – – Anelsonia J.F. Macbr. 11 7 n =7 & Payson Aphragmus Andrz. ex DC. 11 2 7, 8 n =7,8 Aplanodes Marais 2 – – – Arabidella (F. Muell.) 61 12 n = 12, 18 O.E. Schulz Arabidopsis (DC.) Heynh. 11 7 5, 8 n = 5, 8, 12, 14, 16; 2n = 10, 16, 20, 22, 26, 28, 32, 48 Arabis L. 118 59 6, 7, 8 n = 6, 7, 8, 12, 14, 16, 21, 32; 2n =8, 12, 14, 16, 18, 21, 24, 28, 30, 32 Arcyosperma O.E. Schulz 1 – – – Armoracia Gaertn., 33 8or16n = 16; 2n =32 B. Mey. & Scherb. Aschersoniodoxa Gilg 3– – – & Muschl. Asperuginoides Rauschert 1 1 8 2n =16 Asta Klotzsch & 21 10 n =10 O.E. Schulz ex O.E. Schulz Atelanthera Hook. f. 1– – – & Thomson Athysanus Greene 2 1 13 2n =26 Aubrieta Adans. 12 7 8 n =8;2n = 16, 32 Aurinia Desv. 9 9 8 n = 8, 16; 2n =16 Baimashania Al-Shehbaz 2 – – – Ballantinia Hook. f. 1– – – ex E.A. Shaw Barbarea R. Br. 29 12 8, 9 n =8;2n = 16, 18 Berteroa DC. 5 4 8 n =8;2n =16 Berteroella O.E. Schulz 1 – – – Biscutella L. 53 35 6, 8, 9 n =6,8,9,18;2n = 12, 16, 18, 36, 54 Bivonaea DC. 1 – – – Blennodia R. Br. 2 1 12 n =12 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database 239

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n)

Boechera A. Lo¨ ve 65 34 6, 7, 8 n = 6, 7, 8, 14, 15, 16, 21; 2n = 12, 14, &D.Lo¨ ve 21, 28 Boreava Jaub. & Spach 2 2 7 2n =14 Bornmuellera Hausskn. 7 2 8 2n =16 Borodinia N. Busch 1 1 7 2n =14 Botschantsevia Nabiev 1 – – – Brassica L. 39 33 7, 8, 9, n = 7, 8, 9, 10, 11, 16, 17, 18, 19, 20; 10, 11 2n = 16, 18, 20, 22, 24, 27, 32, 34, 36, 38, 40, 44 Braya Sternb. & Hoppe 13 10 14, 16 n = 14, 16, 21, 28, 32, 35; 2n = 28, 32, 42, 48, 56, 64, 70 Brayopsis Gilg & Muschl. 6 – – – Brossardia Boiss. 1 1 14 n = 14; 2n =28 Bunias L. 3 2 7 n =7;2n = 14, 28, 42 Cakile Mill. 6 6 9 n = 9, 18; 2n = 18, 36 Calepina Adans. 1 1 7 n = 14, 21; 2n = 14, 28 Callothlaspi F.K. Mey. 1 – – – Calymmatium O.E. Schulz 2 – – – Camelina Crantz 11 5 6, 8, 10, 13 n = 6, 7, 8, 12, 13, 14, 20; 2n = 12, 16, 26, 32, 40 Camelinopsis A.G. Mill. 1 1 7 n =7 Capsella Medik. 5 3 8 n = 8, 16; 2n = 16, 32 Cardamine L. 197 85 6, 7, 8, 10 n = 7, 8, 12, 14, 15, 16, 20, 21, 24, 26, 26–28, 27, 28, 30, 32, 36, 40, 48, 56, 64, 72, 128; 2n = 14, 16, 18, 20, 24, 28, 30, 32, 36, 40, 42, 42–48, 48, 52–54, 56, 64, 72, 72–88, 80, 88, 96, 110, 112, 116, 119, 128, 130–134, ±208, ±240 Carinavalva Ising 1 – – – Carrichtera DC. 1 1 8 n =8;2n = 16, 32 Catadysia O.E. Schulz 1 – – – Catenulina Soja´k1––– Catolobus (C.A. Mey.) 11 7,82n = 16, 21 Al-Shehbaz Caulanthus S. Watson 17 15 10, 12, 14 n = 10, 12, 14; 2n = 24, 28, 38, 48 Ceratocnemum 11 8 n =8 Coss. & Balansa Chalcanthus Boiss. 1 1 7 n =7;2n =14 Chamira Thunb. 1 – – – Chartoloma Bunge 1 – – – Chaunanthus O.E. Schulz 3 – – – Chilocardamum O.E. Schulz 1 – – – Chlorocrambe Rydb. 1 – – – Chorispora R. Br. ex DC. 11 4 7, 9 n = 7, 9, 14; 2n =14 Christolea Cambess. 21 7 2n =14 ex Jacquem. 240 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Chrysochamela (Fenzl) Boiss. 3 1 8 2n =16 Cithareloma Bunge 3 1 13 n = 13; 2n =26 Clastopus Bunge ex Boiss. 2 2 8 2n =16 Clausia Korn.-Trotzky 6 1 7 2n = 14, 28 Clypeola L. 9 4 7, 8 n = 7, 8, 14, 16; 2n = 14, 16, 32 Cochlearia L. 19 14 6, 7 n = 6, 7, 12, 14, 21, 24; 2n = 12, 14, 18, 24, 26, 28, 36, 38, 42, 48, 54, 60 Coincya Porta & Rigo ex Rouy 6 6 12 n = 12, 24; 2n = 24, 48 Coluteocarpus Boiss. 1 1 7 2n =14 Conringia Heist. ex Fabr. 6 5 7, 9 n =7,9;2n = 14, 18, 28 Cordylocarpus Desf. 1 1 8 2n =16 Crambe L. 34 20 15 n = 15, 30, 45, 60; 2n = 30, 60, 90, 120, 150 Crambella Maire 1 1 11 n =11 Cremolobus DC. 8 – – – Crucihimalaya Al-Shehbaz, 9 5 7, 8, 9 n =8,9;2n = 14, 16 O’Kane & R.A. Price Cryptospora Kar. & Kir. 3 1 7 2n =14 Cuphonotus O.E. Schulz 2 – – – Cusickiella Rollins 2 – – – Cymatocarpus O.E. Schulz 3 – – – Cyphocardamum Hedge 1 – – – Dactylocardamum Al-Shehbaz 1 – – – Degenia Hayek 1 1 8 2n =16 Delpinophytum Speg. 1 – – – Descurainia Webb & Berthel. 48 21 7 n = 7, 14, 21; 2n = 14, 21, 28, 36, 42 Desideria Pamp. 12 2 7 2n =14 Diceratella Boiss. 11 1 11 n =11 Dichasianthus Ovcz. & Yunussov 1 – – – Dictyophragmus O.E. Schulz 2 – – – Didesmus Desv. 2 2 8 n =8;2n =16 Didymophysa Boiss. 2 1 7, 8 2n = 14, 16 Dielsiocharis O.E. Schulz 2 1 7, 8 n =8,14 Dilophia Thomson 2 – – – Dimorphocarpa Rollins 4 3 9 n =9;2n =18 Diplotaxis DC. 32 22 7, 8, 9, 10, n = 7, 8, 9, 10, 11, 13, 21; 2n = 14, 16, 11, 13 18, 20, 22, 26, 38, 42 Dipoma Franch. 1 – – – Diptychocarpus Trautv. 1 1 7 n =7;2n =14 Dithyrea Harv. 2 2 10 n = 10, 30, 40 Dontostemon Andrz. 11 5 7 n =7;2n = 14, 28 ex C.A. Mey. Douepea Cambess. 21 8 n =16 ex Jacquem. S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database 241

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Draba L. 363 138 6, 7, 8, 9, n = 7, 8, 9, 10, 12, 13, 14, 15, 16, 18, 19, 10, 12, 15 20, 24, 27, 28, 32, 36, 38, 40, 48, 56, 64, 72; 2n = 12, 16, 20, 24, 28, 30, 32, 36, 40, 42, 48, 52, 54, 56, 60, 64, 72, 76, 80, 96, 100, 112, 120, 128 Drabastrum (F. Muell.) 1– – – O.E. Schulz Dryopetalon A. Gray 5 2 12, 17 n = 12, 17 Eigia Soja´k1––– Elburzia Hedge 1 1 7 n =7 Enarthrocarpus Labill. 5 5 10 n = 10; 2n =20 Englerocharis Muschl. 2 – – – Eremobium Boiss. 1 1 8, 10, 13 n = 10, 13; 2n = 16, 20 Eremoblastus Botsch. 1 – – – Eremodraba O.E. Schulz 2 – – – Eremophyton Be´g. 1 – – – Eruca Mill. 4 3 11 n = 11; 2n =22 Erucaria Gaertn. 10 7 6, 7, 8 n =6,7,8;2n =16 Erucastrum (DC.) C. Presl 25 17 7, 8, 9 n = 7, 8, 9, 14, 15, 16, 21; 2n = 10, 14, 16, 18, 24, 28, 30, 32, 48, 64 Erysimum L. 223 110 6, 7, 8, 9, 10, n = 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 11, 12, 13, 14, 16, 17, 18, 20, 21, 24, 27, 28; 2n = 12, 15, 16, 17 14, 16, 18, 20, 22, 24, 26, 28, 30, 32, 34, 36, 38, 40, 42, 48, 54, 56, 72 Euclidium R. Br. 1 1 7 n =7;2n =14 Eudema Humb. & Bonpl. 6 – – – Eunomia DC. 1 1 7 n =7;2n =14 Eurycarpus Botsch. 2 – – – Eutrema R. Br. 26 8 7, 9 n = 14; 2n = 18, 28, 36, 42, 56 Farsetia Turra 27 7 6 n = 12, 24; 2n = 12, 24, 60, 72 Fezia Pit. ex Batt. 1 1 11 n =11 Fibigia Medik. 13 6 7, 8 n = 7, 8, 16; 2n =16 Foleyola Maire 1 1 16 n =16 Fortuynia Shuttlw. ex Boiss. 2 1 16 2n =32 Fourraea Greuter & Burdet 1 1 7 2n =14 Gagria M. Kra´l1––– Galitzkya V.V. Botschantz. 3 2 8 2n =16 Geococcus J.L. Drumm. 1– – – ex Harv. Glastaria Boiss. 1 1 7 2n = 14±2 Goldbachia DC. 5 1 14 n = 14; 2n = 14, 28 Graellsia Boiss. 8 3 7, 15 n = 7, 14, 15; 2n = 14, 30 Grammosperma O.E. Schulz 1 – – – Guillenia Greene 1 – – – Guiraoa Coss. 1 1 9 n =9 Gynophora Gilli 1 – – – 242 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Halimolobos Tausch 16 5 7, 8 n = 8, 16; 2n = 14, 32 Harmsiodoxa O.E. Schulz 3 – – – Heldreichia Boiss. 4 – – – Heliophila L. 82 4 10 n = 10; 2n =20 Hemicrambe Webb 3 1 9 n =9 Hemilophia Franch. 5 – – – Henophyton Coss. & Durieu 2 1 42 n =42 Hesperidanthus 51 11 n = 11, 20+ (B.L. Rob.) Rydb. Hesperis L. 46 14 6, 7, 8, 10 n = 6, 7, 8, 12, 14, 16, 20, 24; 2n = 12, 14, 16, 24, 28 Hollermayera O.E. Schulz 1 – – – Hormathophylla Cullen 10 10 8, 11, 14, 15 n = 8, 11, 14, 15, 16, 28; 2n = 16, & T.R. Dudley 22, 28, 30, 32 Hornungia Rchb. 3 3 6 n =6;2n = 12, 24 Horwoodia Turrill 1 1 13 n = 13; 2n =26 Hutchinsia R. Br. 1 – – – Ianhedgea Al-Shehbaz 1 1 14 2n =28 & O’Kane Iberis L. 27 22 7, 8, 9, 11 n = 7, 8, 9, 11, 14, 17, 33–35; 2n = 11, 14, 16, 18, 22, 44, 50ca, 66 Idahoa A. Nelson 11 8 n =8;2n =16 & J.F. Macbr. Iodanthus Torr. & 1– – – A. Gray ex Steud. Ionopsidium (DC.) Rchb. 5 3 11, 12, 16 n = 11, 12, 16; 2n = 22, 24, 32, 36 Irenepharsus Hewson 3 – – – Isatis L. 79 20 7 n = 7, 14, 21, 28; 2n = 14, 28, 42 Iskandera N. Busch 2 – – – Ivania O.E. Schulz 1 – – – Kernera Medik. 1 1 7, 8 n =8;2n = 14, 16 Kremeriella Maire 1 1 12 n =12 Lachnocapsa Balf. f. 1 – – – Lachnoloma Bunge 1 1 7 2n =14 Leavenworthia Torr. 8 8 11, 15 n = 11, 15, 24; 2n = 22, 30 Leiospora (C.A. Mey.) 62 7 2n =14 F. Dvora´k Lepidium L. 231 60 4, 8 n = 4, 8, 12, 16, 32, 64; 2n = 16, 24, 28, 32, 40, 48, 64, 80 Lepidostemon Hook. f. 6– – – & Thomson Leptaleum DC. 1 1 7 n =7;2n =14 Lithodraba Boelcke 1 1 32 2n =64 Litwinowia Woronow 1 1 7 n =7;2n =14 Lobularia Desv. 4 4 6, 11, 12 n = 11, 12, 21; 2n = 22, 24, 36, 42, 48, 96 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database 243

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Lunaria L. 3 2 14, 15 n = 14, 15; 2n = 28, 30, 60 Lyrocarpa Hook. & Harv. 3 2 10 n = 20; 2n = 20, 40 Macropodium R. Br. 2 1 15 n = 15; 2n =30 Malcolmia R. Br. 32 15 7, 8, 10 n = 7, 8, 10, 14, 16; 2n = 14, 16, 20, 24, 28, 32 Mancoa Wedd. 11 1 8 2n =16 Maresia Pomel 3 1 14 2n =28 Mathewsia Hook. & Arn. 10 – – – Matthiola R. Br. 48 24 5, 6, 7, 8, 13 n = 6, 7, 8, 13; 2n = 10, 11, 12, 14, 16, 24, 28 Megacarpaea DC. 9 1 9 2n =18 Megadenia Maxim. 1 1 6, 10 2n = 12, 20 Menkea Lehm. 6 – – – Menonvillea DC. 24 1 11 2n =22 Microlepidium F. Muell. 2 – – – Microstigma Trautv. 3 1 6 2n =12 Microthlaspi F.K. Mey. 4 4 7 n = 7, 14, 21, 35; 2n = 14, 28, 42 Morettia DC. 3 1 11 n =11 Moricandia DC. 8 8 11, 14 n = 11, 14; 2n = 22, 28, 56, 84 Moriera Boiss. 1 1 12 n =12 Morisia J. Gay 1 1 7 2n =14 Murbeckiella Rothm. 5 2 8 n =8;2n =16 Muricaria Desv. 1 1 12 2n =24 Myagrum L. 1 1 7 n =7;2n =14 Nasturtiopsis Boiss. 1 1 7 n =7 Nasturtium R. Br. 10 3 7, 8 n = 7, 8, 14, 16, 32; 2n = 32, 64 Neotchihatchewia Rauschert 1 – – – Neotorularia Hedge 11 4 7 n =7;2n = 14, 28, 42 &J.Le´onard Nerisyrenia Greene 8 7 7, 9, 10, 11 n = 7, 8, 9, 10, 11, 16, 18, 20, 30, 36; 2n = 18, 19+, 20, 21+, 22+, 23, 27, 32, 33, 34, 34+, 36, 38, 40, 41+, 44+, 48, 58, 59+ Neslia Desv. 1 1 7 n = 7, 14, 21; 2n = 14, 42 Neuontobotrys O.E. Schulz 5 – – – Nevada N.H. Holmgren 1 1 7 n =7;2n =14 Noccaea Moench 77 33 7 n = 7, 14, 28; 2n = 14, 28, 42, 56 Noccidium F.K. Mey. 1 – – – Notoceras R. Br. 1 1 11 n = 11; 2n =22 Notothlaspi Hook. f. 2 1 19 n =19 Ochthodium DC. 1 – – – Octoceras Bunge 1 1 7 2n =14 Olimarabidopsis Al-Shehbaz, 32 8 n = 8, 16; 2n = 32, 48 O’Kane & R.A. Price Onuris Phil. 6 1 9 2n =18 Oreoblastus Suslova 1 – – – 244 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Oreoloma Botsch. 3 – – – Oreophyton O.E. Schulz 1 1 16 2n =32 Ornithocarpa Rose 2 1 24 n = 24; 2n =48 Orychophragmus Bunge 2 1 12 n = 12; 2n =24 Otocarpus Durieu 1 1 8 2n =16 Pachycladon Hook. f. 10 1 10 2n =20 Pachymitus O.E. Schulz 1 – – – Pachyneurum Bunge 1 – – – Pachyphragma (DC.) Rchb. 1 – – – Pachypterygium Bunge 3 2 7 n = 14; 2n =14 Parlatoria Boiss. 2 2 7 n =7 Parodiodoxa O.E. Schulz 1 – – – Parolinia Webb 5 4 11 2n =22 Parrya R. Br. 34 3 7 2n = 14, 21, 28 Paysonia O’Kane 8 8 7, 8, 9 n =7,8,9;2n = 14, 16, 18 & Al-Shehbaz Pegaeophyton Hayek 7– – – & Hand.-Mazz. Peltaria Jacq. 4 3 7 2n = 14, 28, 56 Peltariopsis (Boiss.) N. Busch 2 – – – Pennellia Nieuwl. 9 2 8 n =8;2n =16 Petrocallis R. Br. 1 1 7 2n =14 Petroravenia Al-Shehbaz 1 – – – Phaeonychium O.E. Schulz 7 – – – Phlebolobium O.E. Schulz 1 – – – Phlegmatospermum O.E. Schulz 4 1 6 n =6 Phoenicaulis Nutt. 1 – – – Physaria (Nutt.) A. Gray 105 59 4, 5, 7, 8, 9, n = 4, 5, 6, 7, 8, 9, 10, 12, 15, 18, 24, 10, 12, 15 30, 36; 2n = 8, 10, 12, 14, 16, 18, 20, 24, 30, 48?, 52, 60, 67–70 Physocardamum Hedge 1 – – – Physoptychis Boiss. 2 1 8 2n =16 Physorhynchus Hook. 2 2 14, 16 n = 16; 2n = 28, 32 Planodes Greene 1 1 8 n =8;2n =16 Polyctenium Greene 2 – – – Polypsecadium O.E. Schulz 3 – – – Pringlea Anderson ex Hook. f. 1 1 12 2n =24 Pseuderucaria (Boiss.) 21 14 n =14 O.E. Schulz Pseudoarabidopsis 11 6 2n =12 Al-Shehbaz, O’Kane & R.A. Price Pseudocamelina (Boiss.) 31 7 n =7;2n =14 N. Busch Pseudoclausia Popov 10 1 7 n =7;2n =14 Pseudofortuynia Hedge 1 1 7 n =7;2n =14 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database 245

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Pseudosempervivum 3– – – (Boiss.) Grossh. Pseudoturritis Al-Shehbaz 1 1 8 n =8;2n =16 Pseudovesicaria (Boiss.) Rupr. 1 1 x = 5 2n =10 Psychine Desf. 1 1 15 n = 15; 2n = 16, 30 Pterygiosperma O.E. Schulz 1 – – – Pugionium Gaertn. 3 – – – Pycnoplinthopsis Jafri 1 – – – Pycnoplinthus O.E. Schulz 1 – – – Quezeliantha H. Scholz 1– – – ex Rauschert Raﬀenaldia Godr. 2 2 7 n = 7, 10, 14; 2n = 14, 28 Raphanoryncha Rollins 1 – – – Raphanus L. 3 2 9 n = 9, 18; 2n =18 Rapistrum Crantz 2 2 8 n =8;2n = 16, 18 Rhammatophyllum O.E. Schulz 10 1 8 2n =16 Rhizobotrya Tausch 1 1 7 2n =14 Ricotia L. 9 3 14 2n =28 Robeschia Hochst. 11 8 n =8 ex O.E. Schulz Rollinsia Al-Shehbaz 1 1 10 n = 10; 2n =20 Romanschulzia O.E. Schulz 13 – – – Rorippa Scop. 86 34 5, 6?, 7, 8 n = 5, 7, 8, 12, 14, 16, 24, 28; 2n = 16, 24, 28, 32, 40, 44, 48 Rytidocarpus Coss. 1 1 14 2n =28 Sameraria Desv. 9 5 7, 8 n =7;2n = 14, 16 Sandbergia Greene 1 – – – Sarcodraba Gilg & Muschl. 3 – – – Savignya DC. 1 1 15 n =15 Scambopus O.E. Schulz 1 – – – Schimpera Hochst. 11 7 n =7;2n =14 & Steud. ex Endl. Schivereckia Andrz. ex DC. 2 2 8 2n =16 Schizopetalon Sims 10 1 9, 10 n = 10; 2n =18 Schouwia DC. 1 1 18 n = 18; 2n =36 Scoliaxon Payson 1 – – – Selenia Nutt. 4 4 7, 12 n = 7, 12, 23, 69 Shangrilaia Al-Shehbaz, 1– – – J.P. Yue & H. Sun Sibara Greene 9 5 8, 14 n = 8, 14; 2n = 16, 26, 28 Sibaropsis S. Boyd & T.S. Ross 1 1 14 2n =28 Sinapidendron Lowe 4 4 9, 10 n = 9, 10; 2n = 18, 20 Sinapis L. 5 5 7, 8, 9, 12 n = 7, 9, 12; 2n = 14, 16, 18, 24 Sisymbrella Spach 2 2 8 n =8;2n = 16, 32 Sisymbriopsis Botsch. & Tzvelev 5 – – – 246 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Sisymbrium L. 95 29 7, 9, 10, 13 n = 7, 9, 13, 14, 21, 28; 2n = 14, 20, 21, 26, 28, 42, 56 Skottbergianthus Boelcke 1 – – – Smelowskia C.A. Mey. 25 13 6, 10, 11, 12 2n = 12, 18, 20, 22, 24, 32, 36 Sobolewskia M.Bieb. 4 1 8 2n =16 Solms-laubachia Muschl. 9 8 7 2n = 14, 28 Sphaerocardamum S. Schauer 8 2 8 n =8 Spirorhynchus Kar. & Kir. 1 1 7 2n =14 Spryginia Popov 7 – – – Stanleya Nutt. 6 3 6, 7 n = 12, 14, 28; 2n = 24, 28, 56 Stenopetalum R. Br. ex DC. 10 7 4, 5 n =4,5;2n =8,10 Sterigmostemum M.Bieb. 7 4 7 n =7;2n =14 Stevenia Adams ex Fisch. 4 2 16 2n =32 Straussiella Hausskn. 1 1 8 n =8 Streptanthella Rydb. 1 1 7 n = 7, 14; 2n = 14, 28 Streptanthus Nutt. 34 27 12, 14 n = 12, 14; 2n = 14, 28, 56 Streptoloma Bunge 2 1 7 n =7,40 Strigosella Boiss. 4 – – – Stubendorﬃa Schrenk. 8– – – ex Fisch. & Ave´-Lall. Subularia L. 2 2 14, 15 n = 15; 2n = 28, 30 Succowia Medik. 1 1 18 n = 16, 18; 2n =36 Synstemon Botsch. 2 – – – Synthlipsis A. Gray 3 1 10 n = 10; 2n =20 Tauscheria Fisch. ex DC. 1 1 7 n =7;2n = 12, 14 Teesdalia R. Br. 3 3 9, 10 n = 18; 2n = 18, 20, 36 Tetracme Bunge 10 2 14 n = 14; 2n = 14, 28 Thelypodiopsis Rydb. 17 4 10, 11 n = 10, 11 Thelypodium Endl. 19 9 13 n =13 Thlaspi L. 55 8 7 n =7;2n = 14, 28, 56, 84 Thysanocarpus Hook. 5 1 7 n = 7, 28; 2n =28 Trachystoma O.E. Schulz 3 3 7, 8 n =7,8;2n =16 Transberingia 11 8 2n = 16, 20 Al-Shehbaz & O’Kane Trichotolinum O.E. Schulz 1 – – – Tropidocarpum Hook. 4 1 8 n =8 Turritis L. 2 2 6, 8, 9 n = 6, 8, 16; 2n = 12, 16, 32 Vania F.K. Mey. 1 – – – Vella L. 7 6 17 n = 17; 2n = 34, 48, 68, 102 Veselskya Opiz 1 – – – Warea Nutt. 4 1 12 n = 12; 2n =24 Weberbauera Gilg & Muschl. 23 1 13 or 26? 2n =52 Werdermannia O.E. Schulz 5 – – – Winklera Regel 3 – – – Xerodraba Skottsb. 7 1 11 2n =22 Yinshania Ma & Y.Z. Zhao 13 8 6, 7 2n = 12, 14, 42, 44 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database 247

Table 1. (Continued) Genus No. No. Base Chromosome Species Species Chromosome No. counted No. (x) (n,2n) Zerdana Boiss. 1 1 7 2n =14 Zilla Forssk. 2 2 16 n = 16; 2n =32 Zuvanda (Dvora´k) Askerov 3 – – – TOTAL No. genera 338 232 TOTAL No. species 3709 1558

known for the family (n = 4) has been found well documented: including e.g. Brassica in two unrelated genera, the Australian Sten- carinata A. Braun, B. juncea (L.) Czern., opetalum and the western North American B. napus L., B. balerica Pers., Diplotaxis Physaria, while the highest number reported muralis (L.) DC., and Erucastrum gallicum (n = 128) is in the eastern North American (Willd.) O.E. Schulz. Cardamine concatenata (Michx.) O. Schwarz Earlier authors, e.g. Mukherjee (1973), and C. diphylla (Michx.) A.W. Wood. believed that aneuploidy and diminution in Both polyploidy and aneuploidy have chromosome size, rather than polyploidy, played an important role in the evolution played an important role in the evolution of of the Brassicaceae (see review paper by the family. But recent genomic studies (see Marhold in the same issue). Nearly 37% of review paper by Lysak in the same issue) have the species are polyploids, and 10% of the confirmed ancient polyploid (paleopolyploid) genera (Armoracia, Braya, Brossardia, Cram- events in the family (at least in the tribe be, Douepea, Foleyola, Fortuynia, Henophy- Brassiceae), suggesting that higher chromo- ton, Ianhedgea, Lithodraba, Lunaria, some numbers may represent the more prim- Macropodium, Maresia, Moricandia, Notothl- itive state and that lower numbers have likely aspi, Oreophyton, Ornithocarpa, Physorhyn- resulted from chromosome fusions. Numerical chus, Pseuderucaria, Psychine, Ricotia, and structural diversification of chromosomes Rytidocarpus, Savignya, Schouwia, Sibaropsis, appear to have evolved independently in sev- Stevenia, Streptanthus, Subularia, Succowia, eral lines. Vella, Weberbauera, and Zilla), appear to be exclusively polyploids (defined as n ‡ 14). References Intraspecific polyploidy (presumably autopo- lyploidy) is reported for many species (e.g. Jaretzky R. (1928) Untersuchungen u¨ ber Chromo- species of Erucastrum, Coincya, Vella, etc.) somen und Phylogenie bei einigen Cruciferen. and would appear to have played an Jahrb. Wiss. Bot. 68: 1–45. important role in the evolution of species Jaretzky R. (1932) Beziehungen zwischen Chromo- complexes. Intraspecific aneuploidy (e.g. somenzahl und Systematik bei den Cruciferen Cardamine pratensis L. and Draba verna Jahrb. Wiss. Bot. 76: 485–527. Manton I. (1932) Introduction to the general L.) is obviously more frequent at higher cytology of the Cruciferae. Ann. Bot. Lond. I., polyploid series and would not appear to be 46: 509–556. of taxonomic significance. Allopolyploidy is Mukherjee P. (1973) Chromosome study as an aid also a common mode of hybrid speciation in in tracing the evolution of Cruciferae. Cytolgia the Brassicaceae, and many examples are 40: 727–734. 248 S. I. Warwick and I. A. Al-Shehbaz: Brassicaceae: Chromosome number index and database

Addresses of the authors: Suzanne I. Warwick K1A 0C6, Canada. Ihsan A. Al-Shehbaz (e-mail: (e-mail: [email protected]) Agriculture and Agri- [email protected]) Missouri Botanical Food Canada, Eastern Cereal and Oilseed Research Garden, P.O. Box 299, St. Louis, Missouri 63166- Centre, Central Experimental Farm, Ottawa, Ontario 0299, U.S.A.