Pl. Syst. Evol. 259: 89–120 (2006) DOI 10.

1007/s00606-006-0415-z

Systematics and phylogeny of the Brassicaceae (Cruciferae): an overview
I. A. Al-Shehbaz1, M. A. Beilstein2, and E. A. Kellogg2
1 2

Missouri Botanical Garden, St. Louis, Missouri, USA Department of Biology, University of Missouri-St. Louis, St. Louis, Missouri, USA

Received November 9, 2005; accepted December 13, 2005 Published online: June 19, 2006 Ó Springer-Verlag 2006

Abstract. A critical review of characters used in the systematics of the Brassicaceae is given, and aspects of the origin, classification, and generic delimitation of the family discussed. Molecular phylogenetic studies of the family were reviewed, and major clades identified. Based on molecular studies, especially from the ndhF chloroplast gene, and careful evaluation of morphology and generic circumscriptions, a new tribal alignment of the Brassicaceae is proposed. In all, 25 tribes are recognized, of which seven (Aethionemeae, Boechereae, Descurainieae, Eutremeae, Halimolobeae, Noccaeeae, and Smelowskieae) are described as new. For each tribe, the center(s) of distribution, morphology, and number of taxa are given. Of the 338 genera currently recognized in the Brassicaceae, about 260 genera (or about 77%) were either assigned or tentatively assigned to the 25 tribes. Some problems relating to various genera and tribes are discussed, and future research developments are briefly covered. Key words: Brassicaceae, characters, origin, classification, generic circumscription, molecular data, major clades, new tribal alignments.

The Brassicaceae (Cruciferae), or mustard family, is a monophyletic group of about 338 genera and some 3709 species distributed worldwide. It includes many economically important ornamental and crop species (veg-

etables or sources of industrial and cooking oils, forage, and condiments), but is perhaps best known for thale cress, Arabidopsis thaliana (L.) Heynh., the model organism of flowering plants. The latter species is currently used in almost every discipline of experimental biology and its completely sequenced genome (The Arabidopsis Genome Initiative, 2000) paved the way to a better understanding of every aspect of plant biology. The morphological diversity, systems of classification, earlier literature, endemism, and distribution of the family are discussed in Hedge (1976), Al-Shehbaz (1984), and Appel and Al-Shehbaz (2003). These aspects will not be repeated here, and the interested reader should consult those works and the references cited therein. For extensive reviews on the molecular phylogenetic studies of the family, Koch (2003), Koch et al. (2003a), MitchellOlds et al. (2005), and Beilstein et al. (2006) should be checked. The present paper addresses the following major aspects of the family: the evaluation of characters and their utilization in infrafamilial classifications, delimitation of genera, molecular data and major subdivisions of the family,

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problematic taxa, and future challenges of research. Characters Numerous studies (e.g. Al-Shehbaz 1984, Price et al. 1994, Appel and Al-Shehbaz 2003, Koch et al. 2003a, Mitchell-Olds et al. 2005) have amply demonstrated that morphological characters in the Brassicaceae are highly homoplasious, making it virtually impossible to utilize them alone in establishing phylogenetic relationships on a family-wide basis or sometimes even within genera (Mummenhoff et al. 1997a). The lack of a robust phylogeny of the family led some recent authors (e.g. Rollins 1993, Appel and Al-Shehbaz 2003) to adopt an alphabetical system in their enumeration of taxa. Fruit morphology and seed embryo type (position of the radicle in relation to cotyledons) have been used almost exclusively in the delimitation of taxa at all taxonomic levels, but particularly at generic and tribal ranks, while floral, vegetative, and trichome characters have often been considered far less significant. However, as shown below, fruit and embryo features can be subject to considerable convergence and therefore are sometimes taxonomically unreliable. For example, diplecolobal cotyledons (i.e. with incumbent cotyledons folded two or more times), thought to be unique to the tribe Heliophileae (Schulz 1936), evolved independently in Lepidium L. s.l. (Hewson 1981, Mummenhoff et al. 2001a). From that type, spiral cotyledons probably evolved in Brachycarpaea DC. (Appel and AlShehbaz 1997, Mummenhoff et al. 2005), a genus recently reduced to synonymy of Heliophila L. (Al-Shehbaz and Mummenhoff 2005). Incumbent and accumbent cotyledons, the most common cotyledonary types in the family, are the least reliable because they occur within numerous genera, including Arabidopsis (DC.) Heynh. and Erysimum L. Unfortunately, little is known about the genetic control of cotyledonary position, and A. thaliana would be the ideal species to study the evolution of this character.

Although the Brassicaceae was once thought to be exclusively stenopalynous (i.e. with uniform pollen) with only tricolpate pollen (Erdtman 1971), preliminary surveys (e.g. Rollins and Banerjee 1979) demonstrated the existence in the New World of several genera with 4–11-colpate pollen. This group with ‘‘polycolpate’’ pollen has subsequently been shown by O’Kane and Al-Shehbaz (2003) to form a monophyletic clade. However, a more comprehensive palynological survey of the family is needed to determine the utility of pollen in taxonomic and phylogenetic studies. In fact, pollen data were shown to be useful in the separation of putatively closely related genera (Rollins 1979, Al-Shehbaz 1989). Despite the conservative floral architecture of the family, there can be enormous variation among related groups or even within genera. For example, Lepidium, Heliophila, Alyssum L., and Streptanthus Nutt. all exhibit tremendous floral diversity that is quite useful in defining lineages and assessing relationships (Mummenhoff et al. 2001a, 2005). Many other genera (e.g. Stenopetalum R.Br., Schizopetalon Sims, Ornithocarpa Rose, Stanleya Nutt., Warea Nutt., Iberis L., to name a few) are readily recognized by their flowers and evidently are monophyletic. However, little attention has been given to the value of floral morphology in establishing monophyletic groups in the vast majority of the family. Finally, trichome morphology, first emphasized by Prantl (1891) but utilized only a little in subsequent studies (e.g. Rollins and Banerjee 1975, 1976), appears to be far more useful in the separation of closely related genera (e.g. Al-Shehbaz et al. 1999) and holds significant promise in the delimitation of monophyletic groups. Although both simple and branched trichomes can be found in most major clades of the family (Beilstein et al. 2006, Bailey pers. com.), the trichome subtypes can be far more valuable. The extensive studies on trichome development in Arabidopsis thaliana (e.g. Schwab et al. 2000, Hu ¨ lskamp 2000, Szymanski et al. 2000, Beilstein and Szymanski 2004) have identified a significant number

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of genes (e.g. ANGUSTIFOLIA, STICHEL, ZWICHEL) responsible for the genetic pathways that control trichome morphology. However, sequence comparisons from such genes are not available for other genera of the family. It is not yet known if sequence data from such genes are useful in phylogenetic studies in the family. Origin and classification Hayek (1911), followed by Schulz (1936) and Janchen (1942), provided the first theory on the origin of the Brassicaceae. They believed in a New World origin of the family from the capparaceous subfamily Cleomoideae through the ‘‘basal’’ mustard tribe Theylopodieae (Stanleyeae). Views of this German school were followed by Al-Shehbaz (1973, 1985a), Hauser and Crovello (1982), and Takhtajan (1997). Indeed, Nuttall (1834) proposed the name Stanleyeae as a distinct family intermediate between the Capparaceae and Cruciferae. It included Stanleya Nutt. and Warea Nutt. By ´ k (1973) proposed an Old contrast, Dvora World origin from the Cleomaceae via the tribe Hesperideae, but his views were not subsequently followed. Molecular studies (Hall et al. 2002, Koch et al. 2003a and references therein, MitchellOlds et al. 2005, Beilstein et al. 2006) have clearly demonstrated that the Brassicaceae evolved in the Old World and is sister to the Cleomaceae, that Aethionema R.Br. is the most ‘‘basal’’ genus in the family, that the Thelypodieae (hereafter Schizopetaleae; see below) is rather advanced, and that the remarkable superficial floral and fruit similarities (e.g. exserted stamens equal in length, linear anthers coiled at dehiscence, dense racemes, linear fruits, sessile stigmas, long gynophores) between members of this tribe, especially Stanleya and Warea (first discovered species of each was originally described as Cleome L.) and the Cleomaceae evolved through convergence. A closer comparison of Aethionema with some members of the Cleomaceae is given in the section on major clades of the family.

Although Schulz’s (1936) classification of the Brassicaceae has been modified and criticized (e.g. Janchen 1942; Al-Shehbaz 1973, 1984), it continued to be the most widely used to the present. However, all of his new suprageneric taxa that first appeared in that work are invalidly published (see Greuter et al. 2000). Regardless of the number of infrafamilial taxa recognized, his and the earlier systems of Prantl (1891) and Hayek (1911) utilized a limited number of characters, many of which have recently been shown to be subject to convergence. As a result, almost all of their major subdivisions of the family have been shown by molecular studies to be polyphyletic and artificial (Price et al. 1994; Warwick and Black 1994, 1997a, 1997b; Koch et al. 1999a, 2000, 2001, 2003a; Zunk et al. 1999; Bailey et al. 2002; O’Kane and Al-Shehbaz 2003; Mitchell-Olds et al. 2005; Warwick and Sauder 2005; Beilstein et al. 2006). A classic example of the artificiality of previous classifications is illustrated by Arabidopsis, Capsella Medik., Neslia Desv., and Arabis pendula L. (now Catolobus (C. A. Mey.) Al-Shehbaz). Schulz (1936) placed these taxa in the tribes Sisymbrieae, Lepidieae, Euclidieae, and Arabideae, respectively, but molecular data (Koch et al. 2001, O’Kane and Al-Shehbaz 2003, Beilstein et al. 2006) clearly demonstrated that the four genera are very closely related and therefore should be placed in the same tribe/clade (see below). Generic delimitations There is a considerable lack of agreement among authors of the past century regarding the number of genera in the family (Table 1). Of those currently recognized, 225 genera (66% of the total) are either monotypic or oligotypic (with 2–4 spp.). The vast majority of these monotypic and oligotypic genera are nested within, and should be united with, the larger ones (see below). Comparative sequence data of rapidly evolving DNA regions of the chloroplast (e.g. ndhF gene) and nucleus (e.g. ITS)

A.g. some of the larger genera (e. Crespo et al. MADS-box. based on their fruit morphologies. 2001a) and Boleum within Vella (Warwick and Black 1994. 2001a. For example. Mummenhoff et al. Sisymbrium L. FRUITFUL. As shown in Arabidopsis (see below). However. (Al-Shehbaz and Price 2001. Fruit development in Arabidopsis thaliana has been reasonably well studied. Cardaria is nested within Lepidium (Mummenhoff 1995. indehiscence as the main criterion for the delimitation of genera.). Silicularia Compton (1 sp. Mummenhoff et al. and Boleum to synonymy of Vella (Warwick and Al-Shehbaz 1998).e. 2002) and Boleum to that of Vella (Warwick and Al-Shehbaz 1998) are fully justified. and it is quite likely that the same holds for the rest of the family. 2004. These findings should caution the use of fruit dehiscence vs. is having dehiscent instead of indehiscent fruits. 2000).. 2005). and each had to be split into several segregates. Sm. Arabis L. Prior to the utilization of molecular data in phylogenetic studies.). and Thlaspeocarpa C. Crespo et al. and as few as one or double mutant genes can be the difference between dehiscent and indehiscent fruits (Liljegren et al. Mey. Polster 2005). it soon became evident that none of them are. Arabis once included about 180 species worldwide (Al-Shehbaz 1988a). Most of the smaller genera can easily be accommodated within larger ones if the morphological variation of their fruit characters are neither overemphasized nor used as the sole basis for generic delimitation.g. AlShehbaz 2003a) and Euzomodendron Coss. A. Therefore. from Vella L. Al-Shehbaz et al. Arabidopsis. Schlechteria Bolus (1 sp. Other examples are the reduction of Twisselmannia Al-Shehbaz and Agallis Phil. Mummenhoff et al.). molecular studies provide ample support to that view. [2006] 338 20 21 indicate that many taxa show remarkable sequence similarities but drastically different fruit morphologies and embryo positions (Warwick and Black 1994. silicle) and dehiscence (Ferrandiz et al. In fact. SHATTERPROOF) are known to alter fruit shape (i. a relatively small number of genes are responsible for significant alterations in fruit shape. differences in fruit morphology would result in erroneous classifications or generic delimitations and would obscure phylogenetic relationships. Enumeration of genera recognized by various authors Hayek [1911] Total genera Synonyms Added genera 231 11 34 Schulz [1936] 351 21 141 Authors To [1984] 369 88 106 Appel and Al-Shehbaz [2003] 337 59 27 Warwick et al. At least six genes have been identified that control fruit dehiscence in this species.) to synonymy of the larger Heliophila (previously 73 spp. 1999.). A classic example is the reduction of the South African Brachycarpaea (1 sp. Ferrandiz 2002. of . and a few genes (e. 2000. drastic bursts of fruit evolution can rapidly take place that are independent of molecular markers or other aspects of morphology. the reduction of Cardaria to synonymy of Lepidium (Al-Shehbaz et al. 2005).. the only difference that distinguished Cardaria Desv. Beilstein et al. a genus within which all these smaller genera are nested (Al-Shehbaz and Mummenhoff 2005.) were considered to be natural.: Systematics and phylogeny of the Brassicaceae Table 1. from Lepidium and Boleum Desv. Therefore. Thlaspi L. Dinneny and Yanofsky 2004). 2006. In fact. In cases like these. 2000.92 I. 2000. Cycloptychis E.). Dinneny and Yanofsky 2004. length/width ratio. Such data suggest that major changes in fruit morphology can occur rather rapidly and independent of other characters. silique vs. (2 spp. to synonymy of Tropidocarpum Hook. (2 spp.

Tropidocarpum. To conclude. First. all except one of which (S. However. Beilstein et al. Al-Shehbaz et al. Mummenhoff et al. and branched trichomes. Noccidium F. 2006). Microthlaspi F. 1996. Molecular data Numerous phylogenetic or genetic studies of the Brassicaceae have used chloroplast restriction fragment length polymorphisms (RFLPs). As delimited by Meyer (2001)Thlaspi s. The data also show that the apparent similarities in fruit morphology are the result of convergence. Mey. Warwick et al.. and Vania F. (2003a) and Koch (2003) for a complete coverage on those and other markers.: Systematics and phylogeny of the Brassicaceae 93 which 80 occur in North America (Rollins 1993). Sequence comparisons of the internal transcribed spacers of the nuclear ribosomal DNA and the 5. restriction site variation of cpDNA. especially fruit types and embryo position.) are restricted to the Old World. b. The genus was delimited solely on the presence of linear latiseptate fruits (flattened parallel to the septum). and Meyer’s (1973. three important points need emphasis regarding the delimitation of genera. However. Molecular data (Koch et al. this character combination evolved independently in at least 25 genera of the family. 2000. linifolium Nutt.K. because of widespread convergence in most morphological characters. molecular data (Mummenhoff and Koch 1994. but molecular studies (Warwick et al.) of Meyer’s segregates. monotypic or oligotypic genera should not be established without prior molecular studies and critical evaluation of morphology. than to the remaining species previously assigned to it. A. ex Fabr. potentially very useful vegetative and floral characters.str. Sisymbrium still includes about 90 species in the checklist accompanying this issue (Warwick et al. and the examples of Heliophila. The New World taxa previously assigned to Sisymbrium belong to an entirely different and morphologically distinct clade recognized by Warwick and Al-Shehbaz (2003) as the Thelypodieae alliance and herein at the tribal rank. 2006) have clearly shown that the ten segregates of Arabis currently recognized (AlShehbaz 2003b. or amplified fragment length polymorphisms (AFLP) fingerprinting. and it is most closely related to Alliaria Heist. major differences in fruit morphology can be misleading. Mey. 1999. For these reasons. Al-Shehbaz 1986). Al-Shehbaz and O’Kane 2002a). Finally. the genus consists of only 11 species (O’Kane and AlShehbaz 1997. 2005a) have shown that the genus consists of about 40 species. Zunk et al. 2002. Mey. and Vella should be a constant reminder about the dangers of making erroneous taxonomic conclusions by overemphasizing fruit morphology at the expense of other. Second. 1999a. the ITS region) have been the most frequently used .. As shown by AlShehbaz (2003b). they were not included in the present survey.K. As for Arabidopsis. Earlier studies maintain Thlaspi as a large genus of over 80 species. Further molecular studies are needed to establish generic reassignments of the New World species of Sisymbrium. 2005) are unrelated to each other and to Arabis s. though they are remarkably similar in fruit morphology and cotyledonary position. 1997a. Similarly.K. Mey.K. The genus was delimited solely on its angustiseptate fruits (flattened at a right angle to the septum) with four or more seeds. accumbent cotyledons.str. which were based largely on seed-coat anatomy. Koch and Mummenhoff 2001) strongly support the recognition of a few (Callothlaspi F. these characters should be used with extreme care in establishing generic boundaries. and as presently delimited. consists of only six species. O’Kane and Al-Shehbaz 2003.8S rRNA gene (collectively. the interested reader should consult Koch et al. were not recognized (Hedge 1976. Sisymbrium was once thought to consist of 90 species distributed both in the Old and New Worlds (Al-Shehbaz 1988b).. Although such studies provided a wealth of information and were consulted. 2005b).I. 1979) 12 segregates.. the approximately 60 species previously assigned to the genus are now placed in several segregate genera (AlShehbaz et al. Noccaea Moench.

As indicated above. 210 to North America.94 I.).). The references used to compile these approximate figures are too voluminous to enlist in this publication. and psbA-trnH intergenic spacers. 1998). Furthermore. Beilstein et al. multilocus comparative studies that involve several chloroplast. 325 for the trnL-F spacer.). with an emphasis on the delimitation of genera. and the need to clone this marker in such cases in order to obtain much more reliable data.Br.).). Of these. Rorippa Scop. about 100 for the trnT/L spacer. the effects of concerted evolution. Erysimum L.). Lepidium (231 spp. arginine decarboxylase gene (Galloway et al. Isatis L. 180 to Asia. and ca. unpubl. However. ca. Hesperis L. Major clades Despite the incomplete molecular knowledge of the Brassicaceae.). and intensive efforts are needed to utilize single.or low-copy nuclear markers in addition to plastidic and mitochondrial ones.). (86 spp. and trnF (coding for phenylalanine). It should be admitted.). Aethionema (56 spp. (79 spp. 150 to Africa. Lo ¨ ve & D. Alyssum (195 spp. trnLtrnF.e. ca. along with other single copy nuclear and chloroplast genes. 440 for trnL intron. (48 spp. 130 for ndhF. These studies have been critical for determining representative taxa of monpophyletic groups that should be included in the larger family-based phylogenetic analyses. over 730 species have been studied for ITS. markers such as the pistillata gene (Bailey and Doyle 1999). nuclear. 60 for matK.).) are not going to be seriously altered in future studies (species numbers follow Warwick et al.) hold promise. Al-Shehbaz et al. Most have focused on the tribal level or below (i. a number of monophyletic clades or alliances have been identified based . unpubl.). 2006) and it is hoped that more researchers utilize it in their studies. phytochromes (Beilstein. (64 spp.). Perhaps what is most urgently needed is targeting key ‘‘diploid’’ species.: Systematics and phylogeny of the Brassicaceae markers in assessing phylogenetic relationships in the Brassicaceae (see below). trnL (coding for leucine). as well as the trnL intron. and fewer species for other markers. especially with small genome size. that represent major clades or larger monophyletic genera in the family and conduct comprehensive. and mitochondrial NADH dehydrogenase intron (Franzke et al. 80 to South America. 60 to Australia and New Zealand. Heliophila (82 spp.). (48 spp.). Consequently. Boechera A. such as Malcolmia R.Br. 2006). and they have not been utilized on a larger scale in phylogenetic studies. however. and Crambe L.). 250 species are native to Europe. About 930 species in 195 genera of Brassicaceae have been surveyed for one or more markers. need critical studies to establish their monophyly.g. 50 for Chs. at least 70% of the species would be retained in their present genera. (34 spp. Sequence data from the coding chloroplast gene ndhF holds good promise for phylogenetic studies in the Brassicaceae (Beilstein et al.. (53 spp. and they can be obtained directly from the first author. that the utility of ITS sequences can be affected by the frequent occurrence of multiple copies of ITS. Cardamine (197 spp.). Matthiola R. The second most frequently used markers are the non-coding regions from three tRNA genes in the large single-copy region of the chloroplast genome. A. (363 spp. Lo ¨ ve (65 spp.).). as well as Arabis and most members of the tribes Brassiceae and Schizopetaleae (see below). (223 spp. Other medium-sized genera. These include the trnT (coding for threonine). Physaria (105 spp. the sampling for each of the above markers covers less than 50% of the genera. Noccaea (77 spp. Draba L. It is practically impossible to study all genera of the family. ca. Few molecular studies (Koch 2003. the groups that would be most affected are the monotypic and oligotypic genera. 2006) have focused on the establishment of a family-wide phylogeny that would identify all major monophyletic clades. and mitochondrial genes. and Streptanthus (34 spp. Furthermore.). We predict that the boundaries of some of the larger and medium-sized genera (e.Br. ca. (34 spp. Descurainia Webb & Berthel. Biscutella L. and trnT-trnL. Parrya R.). groups of closely related genera). (32 spp. ca.

). (2006). unequal. pers. 1. (2005). such as heterocarpy (see below). Most of these major clades or alliances are recognized herein at the tribal level. Warwick and Sauder (2005). com. and Beilstein et al. The ‘‘basal’’ position of Aethionema does not necessarily mean that it has remained primitive.I.).: Systematics and phylogeny of the Brassicaceae 95 on Kropf (2002). but extensive additional details of each are provided below. (2003a) and Mitchell-Olds et al. as well as mitochondrial markers (Franzke and Mummenhoff. 2005a. com. On the basis of all markers surveyed thus far. Although some aspects of the genus. 2006). Al-Shehbaz et al. The remainder of the family falls into an unresolved polytomy of several major clades within some of which are subclades supported by moderate to high bootstrap values (Fig. are not found elsewhere in the family. 1). It has been estimated that the Brassicaceae probably appeared some 50 million years ago (MYA) and that the split between Aethionema and the rest of the family was about 40 MYA (Koch et al. extremely well-resolved groups. Schranz. including chloroplast and nuclear markers (see Kropf 2002. The next step will be to determine the morphology of the Fig. 2004b. A. The Aethionemeae clade is separated from the rest of the family by 100% bootstrap values (and a substantial branch length). 2004a. It is highly likely that the genus underwent adaptive radiation. we hardly know anything about the evolution of that character. pers. This lack of resolution suggests that major adaptive radiation(s) took place in the early evolutionary history of the family. (2002. Phylogenetic relationships among tribes of the Brassicaceae (modified from Beilstein et al. Beilstein et al. Warwick et al. 2006) . Koch 2003. similar to that observed in the rest of the family. It would be highly desirable to determine as accurately as possible the age of that initial radiation and to have a better understanding of the conditions that prompted it. Some were briefly covered in Koch et al. 2005b). the Brassicaceae are split into two. 2003a. This lack of resolution among the basal portion of the family appears to be the case with every marker surveyed thus far. Koch (2003).

(Cleomaceae). Some species of Aethionema are superficially very similar to the Middle Eastern Dipterygium Decne. Tribe Aethionemeae. alliance. Thlaspi. assemblage... group. fruit morphology and heterocarpy (indehiscent 1seeded samaras or dehiscent. Genera of the 25 tribes listed below were compared to that of de Candolle (1821a. and Noccaea. (NE Africa and Arabia eastward into Pakistan) is a herb or shrub with entire linear leaves jointed at their attachment to the stem and with indehiscent. the two genera are placed in one tribe. Koch 2003. etc. Each tribe is defined morphologically and cytologically. the topology of some of the tribes might shift. but these appear to . Hayek (1911) and Schulz’s (1936) tribal classification to determine if any significant similarity patterns exist. subdivision. but the overall infrastructure and principal component genera would most likely remain unchanged. the topography and resolution among the clades represented by the above genera depended on the markers used. 12. 22.) is centered in Turkey but with fewer species extending eastward into Turkmenistan and westward into Spain and Morocco.Br. along with Thlaspi and Ionopsidium (DC. The family polytomy includes several monophyletic clades represented by the genera Arabidopsis. clade. and the recognition of additional tribes.) Rchb. floral structure (with or without appendages) and color. Prantl (1891). Because of their ‘‘basal’’ and isolated position with respect to the rest of the Brassicaceae (Zunk et al. the placement of further genera. 1980). It is possible that future studies would necessitate revision(s) of the tribal architecture of the family. are given. to enable direct and precise references to them in future studies. (2006). With a better sampling and further molecular studies on the family. Draba. A. 18. 1999. Aethionema shows tremendous variation in habit (annual herbs to shrubs). One might argue that it is premature to propose an incomplete tribal classification of the family because not all genera have been surveyed. Brassica L. Eutrema R. Numbers are based on the comprehensive species checklist (Warwick et al. and base chromosome numbers (n=7. and to avoid the usage of potentially misleading words (e. but molecular data show that Aethionema is not closely related to these two genera. The tribe consists of two genera and 57 species distributed primarily in the Middle East and eastern Europe. sensu stricto. 1996. 14. Taxonomic recognition at the tribal rank is given herein only to the major clades of the family. if any. but it is most likely that the genus is a spiny Aethionema. However. lineage. Schulz (1936) placed Aethionema. He placed Moriera in the subtribe Iberidinae Hayek. 1–8-seeded silicles on the same plant). The overall topology of the family (minus Aethionema) shows major similarities in the analyses of multiple molecular markers by Koch (2003) and ndhF by Beilstein et al. complex.g. Moriera occurs in Afghanistan. a monotypic genus that fluctuated between the Brassicaceae and Capparaceae (Hedge et al. b). Beilstein et al. The main reasons for giving taxonomic ranks to the principal clades are to provide a workable framework for the entire family. several species of Aethionema have the same combination of characters. Indeed. Dipterygium glaucum Decne. 2006). but the tribes proposed herein are much needed to encapsulate the progress made to date and to provide a general framework for future studies. infrafamilial taxon. 2006) and chromosome number index (Warwick and AlShehbaz 2006) provided in this issue.). 16. Al-Shehbaz et al. Aethionema (56 spp. 24.. 11. Hesperis. in the tribe Lepidieae subtribe Thlaspidinae Hayek. or the finer delimitation of the boundaries of each of these tribes would have to rely on future molecular studies. 8. sensu lato.96 I. 30). Iran and Turkmenistan. are symplesiomorphic (shared primitive ancestral characters) in the genus. 21. and the approximate numbers of its component genera and species. as well as its overall distribution range. 1-seeded samaras (Kers 2003).: Systematics and phylogeny of the Brassicaceae ancestral Brassicaceae prior to the Aethionema split from the rest of the family and what characters.

). 2002). as Dipterygium is not basal in the Cleomaceae (Hall et al. This new tribe is almost exclusively North American. and Adh analyses. Heenan and Mitchell 2003. 2001. Price (1 sp. O’Kane et al.Br. chromosome numbers.I. 20 spp.A.) to synonymy of Aethionema. but these were placed in different tribes and subtribes by Hayek (1911) and Schulz (1936).). we know rather little about the genome size. Schulz and Ischnocarpus O. De Candolle (1821a) placed Camelina. However.). Koch et al. all members of the tribe have either terete. Turritis L. Price (9 spp.E.g.). the last three genera belong to the Halimolobeae. Capsella Medik.). These include the New World Sphaerocardamum Shauer. Transberingia Al-Shehbaz & O’Kane (1 sp. Kers (2003) suggested that Dipterygium may be treated as a monotypic family. 1999. the group received less than 50% support in Koch’s (2003) combined ITS. and shows a continuous series x=6–11 in Erysimum. Pseudoarabidopsis AlShehbaz. latiseptate fruits (Al-Shehbaz 2005). Halimolobos Tausch. Schulz (1924. Mey. 1936) placed Arabidopsis. 2002. the systematic position of Eunomia needs to be resolved. the nearest relative of Capsella appears to be Catolobus.) Al-Shehbaz. (including Cheesmania O. Tribe Boechereae. Except for Capsella. Chs. com.).E. 1999a.) showed that Eunomia oppositifolia (Pers. Capsella has angustiseptate fruits and. ex DC.A. Neslia. and several Australian and Asian members. A. However. in the Sisymbrieae subtribe Arabidopsidinae. Because of extensive use of Arabidopsis thaliana in experimental biology. together with a heterogeneous assemblage of genera.f. Although this tribe and the next eight are related and. The base chromosome number is predominantly x=8. (1 sp. matK.). Beilstein et al. Pachycladon Hook. the genus and related genera have been well studied (e. Al-Shehbaz and O’Kane 2002a. Schulz) (10 spp. is unrelated to Aethionema or Iberis and should therefore be re-instated as independent genus. Price et al. The Boechereae includes seven genera and about 110 species most of . O’Kane & R. but the placement of the Australian and some Asian genera remain uncertain. 1997. Boechera furcata (Turcz. form a clade with 78% bootstrap support in the ndhF phylogeny of Beilstein et al. O’Kane & R. Olimarabidopsis Al-Shehbaz. and perhaps Stenopetalum R. a genus with linear.). or quadrangular fruits. As shown below.) Al-Shehbaz (1 sp. 1994. though it is reduced to n=5 in Arabidopsis thaliana and n=4 in Stenopetalum (Shaw 1972). they have silicles instead of siliques. each). 1995. The Camelineae includes primarily annuals (most Erysimum are perennials) with stalked or sessile stellate trichomes often mixed with simple ones (Erysimum has exclusively sessile stellate or malpighiaceous trichomes). pers.).A. Al-Shehbaz et al. Price (3 spp.). Erysimum (223 spp. but Hall et al. Despite the important role that Aethionema holds in understanding the early evolution of the family. (1 sp. together with Neslia and Camelina. and Stenopetalum in the Camelineae. Crucihimalaya AlShehbaz. with minor representations in North America and Australia-New Zealand (ca. com. a tribe related to the Camelineae. Tribe Camelineae. Al-Shehbaz et al. Mummenhoff and Hurka 1994. (2 spp. 2001. as does the taxonomic placement of the ca. and what makes it most basal in the family. (10 spp. latiseptate. 2000. It includes Arabidopsis (11 spp.).) DC. and thus far only one species. (5 spp.A. together. its phylogeny. Mitchell and Heenan 2000. Heenan et al. Appel and Al-Shehbaz (2003) reduced Eunomia DC.). (2006). Camelina Crantz (11 spp.: Systematics and phylogeny of the Brassicaceae 97 be the result of convergence. 2006). 2003. The tribe includes 12– 13 genera and some 240 species distributed primarily in Eurasia.). Neslia Desv. 16 species previously assigned to it. These molecular studies strongly support the close relationship among these genera. O’Kane and Al-Shehbaz 1997. grows in the Russian Far East (Al-Shehbaz 2005). and morphology of its basal species.). (2002) and Menke (pers. Phylogenetic studies addressing all of these matters are in progress (Menke. Catolobos (C. Pennellia Nieuwl.. O’Kane & R.

and Sandbergia Greene . All members of the tribe typically have a base chromosome number of x=7. (11 spp. It is an exclusively New World group. and branched trichomes (absent or in few Boechera and simple in Nevada).: Systematics and phylogeny of the Brassicaceae which belong to Boechera. and the ditypic Cusickiella Rollins and Polyctenium Greene. white (rarely purplish flowers). Lyrocarpa. and Capsellinae (Nerisyrenia (as Greggia A.). Anelsonia. in prep. Lesquerella. its members were placed by Schulz (1936) in various tribes. Holmgren. as well as the ndhF studies by Beilstein et al. The Physarieae are readily distinguished from the rest of the Brassicaceae by having pollen with four or more colpi (the rest of . is circumpolar (Al-Shehbaz and O’Kane 2002b. though three reach the southwestern United States and several are disjunct in South America (Bailey 2001.). 150 species. Physaria was placed in the Lepidieae subtribe Physariinae. and Synthlipsis were originally assigned to the Physarieae by Robinson (1895) and are retained here. consists of seven genera and ca. Gray).E. arctica (Wormsk. and Boechera (as Arabis) in the tribe Arabideae. (2006) and Bailey et al. In the combined molecular and morphological analysis of Bailey et al.F. first recognized by Bailey et al. whereas both Mancoa and Cibotarium O. and much of its complexity is the product of hybridization. Iberidinae (Dithyrea). Pennellia. (2002) as the halimolobine alliance. Tribe Halimolobeae. Pennellia (9 spp.). ebracteate racemes (except in two Mancoa). whereas Lesquerella (now a synonym of Physaria. A. Tribe Physarieae. The genera Dithyrea. O’Kane and Al-Shehbaz 2004). The tribe includes Halimolobos (7 spp. This new tribe. Although the tribe Physarieae is welldefined. Physaria. Phoenicaulis Nutt. and Sphaerocardamum in the Sisymbrieae subtribe Arabidopsidinae. Phoenicaulis. the tribe formed a monophyletic group. (pers. Al-Shehbaz et al. (2006). & Harv. and placed Cusickiella (as Cusickia A. a Mexican-southwestern US center and a South American center. ex Hornem) O’Kane & Al-Shehbaz. The tribe. Nerisyrenia Greene (8 spp. P. (3 spp. & Payson. reduced Polyctenium to Smelowskia of that tribe. com. In addition to the approximately 105 species of Physaria (including Lesquerella S.). The majority are perennials with a well-defined basal rosette. and only five species of Physaria are disjunct in South America (N Argentina and S Bolivia) and one.). Sphaerocardamum (8 spp. Paysonia O’Kane & Al-Shehbaz (8 spp. It is distributed primarily in North America.).). mostly entire leaves (except Polyctenium and Sandbergia). (2 spp. but extensive molecular studies (summarized in Al-Shehbaz 2003b) indicate that the two genera are unrelated. includes five genera and about 40 species. seeds mucilaginous when wetted. Lyrocarpa Hook. Bailey 2001). For example. Schulz (1936) recognized Sandbergia as a member of the tribe Sisymbrieae. and a new genus of eight species to be segregated from Halimolobos (Bailey and Al-Shehbaz. 1936) placed Halimolobos.H. Schulz (1924. Both Mancoa and Pennellia have disjunct centers of distribution. Nevada N. often spreading sepals. Gray) and Synthlipsis). and a base number of x=8. The Halimolobeae have branched trichomes. Monophyly of the tribe is supported by the works of Beilstein et al. see Al-Shehbaz and O’Kane 2002b) was placed in the Drabeae. first identified as a monophyletic clade by O’Kane and AlShehbaz (2003). the ranges of most species of which are in central and northern Mexico.). and the rest represent the monotypic Anelsonia J. and apomixis (see below). the tribe includes Dimorphocarpa Rollins (4 spp. Fuentes-Soriano 2004). Rollins (1993) treated all species of Boechera as members of Arabis. and Synthlipsis A. Schulz were assigned to the Lepidieae because of their angustiseptate silicles.. polyploidy. Wats. Dithyrea Harv. Boechera is taxonomically quite difficult.). Cibotarium and Sphaerocardamum were shown to be congeneric (Rollins 1984.). Mancoa Wedd.). He placed the remaining genera in the Lepidieae subtribes Lyrocarpinae (Lyrocarpa).). Macbr.98 I. Gray (2 spp. (2002).).

(2002a. Selenia in the Lunarieae. Synthlipsis. have been reported. Obviously. Ornithocarpa (2 spp.Br. & Scherb. (2 spp. Franzke et al. Aneuploid series above and below x=8.). simple or no trichomes. Physaria. the two are sufficiently distinct. the Eurasian Armoracia P.). A.). especially in Cardamine (see Marhold and Lihova in this issue) and Rorippa. Nasturtium R. two or more ovules per locule. and angustiseptate or inflated silicles (some Nerisyrenia have siliques). Members of the Cardamineae grow predominantly in mesic or aquatic habitats and are characterized by having pinnately divided or compound (rarely palmately compound or simple) leaves. (8 spp. (1999. and Nasturtium is closest to Cardamine. Leavenworthia Torr. Iberis. and a base number of x=8..).) O.) is closer to members of the Cardamineae than to those of other tribes. Ornithocarpa in the Schizopetaleae. Gaertn. one ovule per locule. latiseptate or terete (angustiseptate in Armoracia) fruits. & Ave (3 spp. and phylogeny of this tribe are currently being studied by Sara FuentesSoriano.). and Hornungia Rchb.).).. Tribe Lepidieae. and a base number of x = 8. 2 are native to Mexico and the United States). Iodanthus in the Matthioleae.) (231 spp. Mey.).I. Subularia L. (1998).. whereas Rorippa is nearest to Barbarea. Most species have a base chromosome number of x=8. Muell. Sweeney and Price (2000). are assigned in this account to at least nine tribes. accumbent cotyledons. Capsella. the monotypic Acanthocardamum Thell. (8 spp. but this assumption needs to be tested with molecular data. (Afghanistan) and Delpinophytum Speg. & A. The tribe has been subjected to several molecular studies [including Les (1994). his delimitation of this tribe. Schulz (6 spp. Based on morphology. though a continuous series of aneuploid reduction to n=4 and increase to n=11. Lyrocarpa. The Lepidieae consists of three to five genera and over 240 species. Cochlearia L. have been well documented in the tribe. angustiseptate fruits evolved independently many times within the family. B. (29 spp. The tribe is characterized by the presence of angustiseptate fruits (secondarily inflated in two species previously placed in Cardaria). and Stroganowia Kar. and Iti Garn. & Kir. Genome size.: Systematics and phylogeny of the Brassicaceae 99 Brassicaceae are tricolpate). (Argentina). (4 spp. It is represented by core genus Lepidium (including Cardaria Desv.E.). forked. Schulz (1936) placed Armoracia in the tribe Drabeae. and Bleeker et al. Although Schulz (1936) placed all of the above genera in the Lepidieae. Tribe Cardamineae. Thlaspi. The artificiality of such circumscription led to his assignment of many unrelated genera to this tribe.) and Winklera Regel Fisch. 2002b).). and Selenia Nutt. both of which grow on all continents except Antarctica. Bleeker et al. as well complex polyploidy. The extensive molecular studies on . and perhaps the Middle Eastern and Central Asian Stubendorffia Schrenk ex ´ -Lall. (3 spp. Franzke et al. and references therein] that support its monophyly. Planodes Greene (1 sp. chromosomal evolution. For example. Al-Shehbaz et al. Barbarea R.-Jones) (197 spp. it appears that the Australian Arabidella (F. simple or no trichomes. might belong to this tribe.. and the remaining five genera in the Arabideae. plus various ploidy levels. which is distributed on all continents except Antarctica.. Tropidocarpum. but as shown by AlShehbaz and Price (1998).. This tribe of ten genera and over 340 species includes the core genera Cardamine (including Dentaria L. typically sessile stellate trichomes (though simple. b).) and Rorippa (86 spp. was based solely on the presence of angustisptate fruits. and the North American Iodanthus Torr. often mucilaginous seeds. He reduced Rorippa to synonymy of Nasturtium. (1 sp. Hedinia Ostenf. Gray ex Steud. which he divided into 13 subtribes. which also occupy aquatic or mesic habitats. (5 spp. (1999). Coronopus Zinn. A reversal to the tricolpate state apparently occurred in one species of Lyrocarpa. and stalked substellate trichomes occur in Paysonia species). Isatis and relatives.Br. Aethionema.).

Farsetia Turra (27 spp. the tribe would probably consist of over 280 species the bulk of which (ca.) are perhaps unrelated and worthy of separate recognition. A group of several Australian species of Lepidium that are shrubs with either incumbent or diplecolobal cotyledons (Hewson 1981) seem to form a distinct group separate from the rest of the genus (Mummenhoff. com. It would be interesting to subject this group to further studies to elucidate its generic and tribal status.).). though closely related. and Goodson (pers.). It is unlikely that Alyssum is monophyletic and that some of its segregates (e. Ptilotrichum C. and the smaller Eurasian Hornungia (3 spp.V. dendritic or rarely only forked trichomes.) have been included in molecular studies. A. Al-Shehbaz et al. diplecolobal cotyledons evolved in this group independently from that of the South African genus Heliophila. Galitzkya V.R. to which it shows more morphological affinities than to Alyssum. 1936) treated the Descurainieae as a subtribe of the Sisymbrieae. predominantly yellow flowers.). Botschantz. appear to merit independent status (see below). usually appendaged filaments. Mummenhoff et al.A. ex O.).). but the available data clearly indicate that the two taxa are remotely related.). A. com. Price (pers.). com. This new tribe consists of about six genera and some 60 species distributed in the Americas.). Mey. Detailed phylogenetic and systematic studies are much needed in order to determine tribal assignment of genera. Alyssum obovatum (C.) Turcz. and a . The Alysseae is characterized by having stellate trichomes. (2 spp.100 I. often numerous tiny seeds.. Schulz and the Patagonian Trichotolinum O. Aurinia Desv. 195) falls in Alyssum.g. but evidently that similarity evolved through convergence.). (2006).).). The tribe is Eurasian and North African.).). and a base number of primarily x=8. as well as the position of the Alysseae in relation to the other tribes. and only one species. Clastopus Bunge ex Boiss.). (2 spp. Asperuginoides Rauschert (1 spp. (4 spp. Clypeola L. (2 spp. North-South American Tropidocarpum (4 spp. (1 sp. Hormathophylla Cullen & T. Tribe Descurainieae. extends it native range from northern and central Asia into northern North America. The Alysseae are maintained herein as a tribe because the vast majority of Alyssum and at least some of the genera would form a monophyletic group that deserve a tribal rank. Bornmuellera Hausskn. central Asian Ianhedgea Al-Shehbaz & O’Kane (1 sp. The tribe consists of herbs (Descurainia is secondarily woody on the Canary Islands). Physoptychis Boiss. and Straussiella Hausskn. (5 spp. Mey. (9 spp. incumbent cotyledons. Descurainia is centered in the Canary Islands (7 spp.). The tribe most likely also includes Alyssoides Mill. with petiolate. and Lobularia Desv.). (13 spp.E.). he included Smelowskia and its allies in that subtribe.). Dudley (10 spp.). and the monotypic Middle Eastern Robeschia Hoschst. Tribe Alysseae. pers. Bowmann et al. The tribe appears to be monophyletic based on preliminary studies by Beilstein et al. (2 spp. mostly few-seeded silicles.: Systematics and phylogeny of the Brassicaceae Lepidium and related genera (Mummenhoff 1995. As delimited by Schulz (1936) and expanded by Dudley and Cullen (1965). Trichome morphology in the tribe shows remarkable similarity to that of the Physarieae.) and North and South America. often winged seeds. 2001a. Didymophysa Boiss.). and Africa. Only a few species each of Alyssum. Eurasia. and members of the two tribes are readily distinguished by the number of pollen colpi discussed above. 2004) were relied on to draw the boundaries of this tribe. (9 spp. Schulz (1924. (7 spp. (3 spp. 1999. Evidently. though lower and higher aneuploid series exist but are rare.E. Berteroa DC. 1–3-pinnatisect cauline leaves. Fibigia Medik. latiseptate or terete (rarely angustiseptate). It includes up to 48 species of Descurainia (including Hugueninia Rchb.). The last two might not be sufficiently distinct from Descurainia. Furthermore. Schulz. Degenia Hayek (1 sp. but the two groups. The latter genus likely belongs to the tribe Arabideae.

Al-Shehbaz 1973. Turritis. may well belong to this tribe. Koch (2003) showed the ‘‘tribe’’ to be sister to Arabis turrita L.Br. but these two genera should probably be assigned to the Arabideae. Guillenia Greene.I. but it is definitely much less than the 118 spp.). The expanded Smelowskia is centered in eastern and central Asia. and Fourraea Greuter & Burdet and others. Sophiopsis O.: Systematics and phylogeny of the Brassicaceae 101 base number of x=7. As delimited by Schulz (1936). On overall morphological grounds. and only seven species are native to northern North America. Tribe Arabideae. Ermania Cham. the tribe consists of perennials (sometimes secondarily annual) with white (rarely cream) flowers. Sinosophiopsis Al-Shehbaz (3 spp. Borodinia N.).A. and Berteroella (1 sp. Sophiopsis. a structure not found elsewhere in the family. (2 spp. & Karav. and Schivereckia Andrz. In addition to the pinnatisect and petiolate cauline leaves and branched trichomes.Busch). Based on the presence of pinnatisect leaves and branched trichomes. Aubrieta (12 spp. Arabis (excluding Boechera. He distinguished the Arabideae solely by the presence of latiseptate . 2002).). (1 sp.. It is likely that the eastern and central Asian Stevenia Adams & Fisch. and Macropodium R. ex Botsch. nonmucilaginous seeds. several. within Smelowskia (Warwick et al. Molecular studies supported the inclusion of Hedinia (4 spp. and perhaps Athysanus Greene (2 spp. Sibara Greene). see recent revision by Al-Shehbaz and Warwick 2006a). Mey. though an aneuploid reduction to x=6 is found in Hornungia. Boechereae (Phoenicaulis.). Kardamoglyphos Schltdl. Barbarea. ex DC. a genus that Schulz (1936) had placed in the Arabideae. now Pseudoturritis Al-Shehbaz (1 sp. he (1936) assigned it to the tribe Lepidieae because of having angustiseptate fruits. ex Botsch. and Gorodkovia Botsch. However.. (Smelowskieae). incumbent cotyledons. A.to many-seeded fruits. often treated as a synonym of Alyssum (Zhou et al. and Dontostemon Andrz.). (Anchonieae). Camelineae (Cheesmania. ex C.) is a monophyletic genus (Koch and Al-Shehbaz 2002). Tribe Smelowskieae..E. The Arabideae comprise at least six genera and over 460 spp. & Schltdl. 1936) placed Smelowskia. and base chromosome number of x=6. Al-Shehbaz et al. Draba (363 spp. The tribe also includes Ermania Cham. and Redowskia in the Sisymbrieae subtribe Descurainiinae. Baimashania AlShehbaz (2 spp. The last genus was assigned by several authors (e. Schizopetaleae (Pleurophragma Rydb.).) Pachyneurum Bunge (1 sp. Schulz (1924.). listed by Warwick et al.) belongs to the Arabideae. Anelsonia. but it is very likely that these characters evolved independently in the two taxa. Despite being the largest in the family and one of the most diversified morphologically.).). glandular papillae. (2004b) demonstrated that there are no solid morphological grounds to maintain the smaller genera above as independent of Smelowskia.). Hauser and Crovello 1982) to the tribe Schizopetaleae (as Thelypodieae) because it has stipitate fruits and exserted stamens. the Arabideae consisted of genera (or their synonyms) herein assigned to the Cardamineae (Cardamine. Erophila DC. Koch. (4 spp. Schulz 1936.). Planodes.). Although Hedinia also has the same leaf and trichome characters.g. and the monotypic Redowskia Cham. distributed primarily in Eurasia and North America north of the Tropic of Cancer (except 70 species of Draba distributed along the high Andes of South American from Colombia southward into Patagonia). Many species of Descurainia have unicellular. Cardaminopsis Hayek). Ptilotrichum. Hayek 1911. see below). whose higher estimate resulted from the lack of thorough phylogenetic study on the genus. This new unigeneric tribe consists of Smelowskia (25 spp.). The exact number of species in Arabis is unknown. though higher polyploids with n=12 and aneuoploid reductions to 11 and 10 have been well documented. Leavenworthia. Eutremeae (Neomartinella Pilg. (2006). Schulz (4 spp. The tribe consists of Draba (including Drabopsis K. Warwick et al. 2004b). Nasturtium.

Lysak et al. both genera should be removed from the Brassiceae. (2006) and discussed above. and/or segmented (heteroarthrocarpic) fruits and simple or no trichomes (for genera and number of ´ mez-Campo 1999. the bulk of his tribe Drabeae consisted of Draba (as delimited herein). native to North America. (1999a. and Pseudoturritis are among its recent generic segregates that are neither closely related to Arabis nor to each other (Al-Shehbaz 2003b. but beginning with the works of Koch et al.102 I.E.g. 1997b. see Go Al-Shehbaz 2003. Warwick and Sauder 2005 and references therein). The delimitation of the tribe has not changed drastically since the detailed work of Schulz (1919. By contrast. Turritis belongs to the Camelineae. Turritis was established in 1753 by Linnaeus. A. Warwick and Sauder 2005. A. Schulz). for example. 2005. this tribe has received considerable molecular studies (summarized in Warwick and Black 1997a. Warwick and Sauder 2005). The tribe consists of 46 ‘‘genera’’ and about 230 spp. No other group in the entire family shows as much fruit diversity as that of the . Fourraea. Beilstein et al.) and Conringia Heist. neither of which has the conduplicate cotyledons or the segmented fruits. Despite the redefinition of its limits. Arabis remains paraphyletic and heterogeneous because its type species (A. alpina L. nonmucilaginous seeds. this genus has been considerably reduced in size. Akeroyd 1993. Arabis consisted of about 180 species (Al-Shehbaz 1988a. Arabis is undoubtedly the most problematic genus in the group. but recent studies (Anderson and Warwick 1999. Catolobus. Tan 2002) unite it with Arabis. Al-Shehbaz et al.) and Pseuderucaria (Boiss. (6 spp. Rollins 1993.str. Tribe Brassiceae. 2002) based solely on superficial morphological grounds. 2000) and subsequently O’Kane and Al-Shehbaz (2003). As shown by Koch (2003) and Beilstein et al. 2005). Except for the four species of Cakile Mill. and Trochiscus O. though higher and lower aneuploidy series and extensive polyploidy are common in Draba. Prior to molecular studies. Because of the economically important Brassica and its relatives.E. and the alleged conduplicate cotyledons present in Calepina and a species of Conringia do not appear to be homologous to those of typical members of the tribe. Calepina Adans.). and a base number of x = 8. (1 sp. the Brassiceae is primarily Mediterranean and southwest Asian. Al-Shehbaz 1985b. and more studies are needed to establish its monophyly and delimit its boundaries. Therefore. FranciscoOrtega 1999. is said to be native to Europe. The Arabideae are characterized by having branched trichomes. (2 spp. and clearly is remotely related to Arabis s. In our opinion. hirsuta (L. and it is an ideal example where convergence in fruit morphology has led to unreliable and chaotic taxonomy (see Al-Shehbaz 2003b). Gomez-Campo 1999).) is sister to Draba and Aubrieta rather than to most of the Eurasian and North American species still retained in it (Koch 2003).f. and North America (Rollins 1993.) Scop. characterized primarily by having conduplicate cotyledons. Schulz (2 spp. The few exceptions to this character combination are Ammosperma Hook.E. Mulligan 1996. often latiseptate fruits (terete in Berteroella and some Draba and Aubrieta Adans. 2006) clearly support their exclusion from this tribe. but most recent authors (e. Asia. the genus remains problematic. Zhou et al. though its range extends southward into South Africa. Lesquerella (now Physaria of the Physarieae). ex Fabr. and would appear to involve more than one species.) O.) were once included in the Brassiceae (Schulz 1936. accumbent cotyledons (incumbent in Berteroella O. Boechera. Rollins 1993). Species delimitation is also problematic. all of which support it as the only monophyletic tribe among the 19 recognized by Schulz (1936). 1923). Cusickia (now Cusickiella of the Boechereae). two character states that evolved independently or together many times in the Brassicaceae.). Appel and species.: Systematics and phylogeny of the Brassicaceae siliques and accumbent cotyledons. entire or dentate leaves. Schulz (now Rorippa) and Armoracia of the Cardamineae.

Tribe Schizopetaleae. Diplotaxis DC. Presl. as tradition´ mez-Campo ally recognized (Schulz 1936. Sinapis L. Dictyophragmus O.g. Catadysia O. form a reasonably well-defined clade with over 90% bootstrap support in Beilstein et al. Lithodraba Boelcke (1 sp. Schulz (2 spp. was discussed above.). Warea Nutt. (4 spp. (23 spp. they will bring the total in the Schizopetaleae to over 330 species and about 35 genera. The vast majority of genera are readily recognizable by their fruits. The Schizopetaleae consist of about 230 species in at least 20 genera. & Bonpl. (6 spp. Eruca Mill.). and therefore obstructed the true relationships within the tribe and led to inadequate taxonomy.. as opposed to the tremendous fruit differentiation. Sarcodraba Gilg & Muschl. Except for a few genera such as Cakile (perhaps including Erucaria Gaertn.). Romanschulzia O. Thelypodium Endl. and Didesmus Desv. (1 sp. have most likely occurred independently of other aspects of morphology. (5 spp. Warwick et al. (3 spp. 1997b.). If added. and some 40 South American species await generic assignments.f.S. Francisco-Ortega 1999.). Hesperidanthus (B. but in every other aspect of vegetative and floral morphology..E. The Brassiceae. Englerocharis Muschl. Hemicrambe Webb.I. Dryopetalon A. and Werdermannia O. Chaunanthus O. and floral differentiations. but the vast majority of botanists believe that taxonomy must reflect phylogenetic data. (34 spp.)..). and Isatideae. (South Indian Ocean islands) are restricted to the New World.). Schulz (5 spp. (2 spp. Schizopetalon (10 spp. It is concluded that rapid evolutionary bursts of fruit morphology.).E. Erucastrum C.E.). 2002 and references therein). Dactylocardamum Al-Shehbaz (1 sp. Stanleya Nutt.).).). and nomenclatural changes will have to be made sooner or later. Hirschfeldia Moench) need to be abandoned despite the fact they are ‘‘traditional’’ and include economically important or weedy taxa.). (9 spp. In addition to this genus. (17 spp. Thysanocarpus Hook. Go 1999). the tribe includes Caulanthus S. Schulz (13 spp.E. & Arn. and Xerodraba Skottsb.) Rydb. Generic limits. Onuris Phil. Vella (including Euzumodendron and Boleum). which are probably controlled by a relatively few genes.).). This lack of molecular.E. & Muschl. (7 spp. (6 spp. Streptanthus Nutt. some of the most commonly known genera of the Brassiceae (e.). Schulz (2 spp. all except the monotypic Pringlea Anderson ex Hook.: Systematics and phylogeny of the Brassicaceae 103 Brassiceae. Gray (5 spp.). This is earliest pub´ oud 1845). The extensive molecular studies by Warwick and her colleagues (see Warwick and Sauder 2005) on this tribe clearly demonstrate that generic boundaries. Rob.). A. The limits of the Schizopetaleae are still unclear. Eremodraba O.). Al-Shehbaz et al.). Schulz (1 sp.). (6 spp. Eudema Humb.).). need to be revised.). (5 spp. Petroravenia Al-Shehbaz (1 sp. Streptanthella Rydb. Thleypodiopsis Rydb. vegetative. Neuontobotrys O. Schulz (2 spp. (19 spp.).). Sibaropsis S. Raphanus L. Sinapidendron Lowe. linifolium) belong to the Schizopetaleae (as Thelypodieae).).). (10 spp.E.L. (2002) demonstrated that all except one of the New World species previously assigned to Sisymbrium (S. and lished name for the tribe (Barne the name should replace the most commonly used Thelypodieae. and Crambe (see Warwick and Black 1994. Sibara Greene.) should be studied in connection with this tribe. and the South American Brayopsis Gilg.. they are inadequately distinguishable. Brassica. Schulz (5 spp. Chlorocrambe Rydb. if indeed possible to establish within the rapa and nigra clades. Rapistrum Crantz. (17 spp. (6 spp.E. Mathewsia Hook. . Sisymbrieae. traditionalists would resist such major alterations.). (1 sp. (2006) but are less resolved in Koch (2003). together with the Schizopetaleae. Boyd & T. Chilocardamum O. Naturally. Schulz (3 spp. should reflect the extensive molecular data available. In fact. Ross (1 sp. and molecular marker surveyed thus far. Weberbauera Gilg & Muschl. the rest of the Brassiceae falls into two groups somewhat weakly defined molecularly but not morphologically: the nigra and rapa clades.).). Wats.).E.

its reduction. Koch 2003). and it is likely that some of Greene’s seven segregates (see Appel and AlShehbaz 2003) would merit recognition. The fruits are primarily siliques. such as Murbeckiella Rothm. inclusion of stamens. Thelypodium and Thleypodiopsis (Hesperideae). 2004b. Floral diversity includes differences in filament length (equal. 2006) or the Brassiceae (Warwick et al. especially in Streptanthus. Al-Shehbaz et al. and about 20 South American genera and over 40 species of Sisymbrium (Sisymbrieae). insect pollination. A. the Schizopetaleae exhibit tremendous floral (instead of fruit) diversity. the base chromosome number is primarily x = 14 and its descending aneuploid series to x = 10. 2006) resolution among members of the Schizopetaleae suggests a relatively recent evolution of the group and insufficient time for the molecular markers studied to diverge. (5 spp. These include Dryopetalon and Schizopetalon (Schizopetaleae). 2002. often urceolate calyx. absence of gynophore. Pringleeae. elaboration of corolla vs. Streptantheae.: Systematics and phylogeny of the Brassicaceae As delimited by Al-Shehbaz (1973). 2004a. species with typically streptanthoid flowers ought to be studied carefully to determine whether they represent one genus or their character combination evolved independently. It would also be a mistake to unite members of the Schizopetaleae into one or few genera. a base . and somewhat zygomorphic flowers). stamens in three lengths. wind vs. but molecular data (see Warwick et al.) and Nasturtiopsis Boiss. Thysanocarpus (Lunarieae). zygomorphy. These acute problems need to be addressed critically by additional molecular and morphological studies. This genus is quite heterogeneous in floral morphology. absence of style. This poor resolution is in agreement with the difficulty in recognizing genera in the tribe. the tribe includes four of Schulz’s (1936) tribes (Stanleyeae. In our opinion.E. It is premature to make rigid conclusions when not all genera have been examined and where molecular data to date provides little support for generic delimitation. belong to this tribe. they were maintained by Rollins (1993) and Appel and Al-Shehbaz (2003) because of small differences in fruit compression and cotyledonary position. Although not all species of the tribe are surveyed cytologically. actinomorphy vs. However. 2002. exsertion vs. and Streptanthus are unique in the family by having this flower combination. presence vs. given the remarkable floral differentiation in the group.). 2002. presence vs. but silicles must have evolved independently in the North American Thysanocarpus and several South American genera. Hesperidanthus (Matthioleae). The tribe is monophyletic and is sister to the Schizopetaleae (Beilstein et al. Unlike the Brassiceae. and every conceivable flower color in the family. or three unequal lengths). Many genera presently assigned to the Schizopetaleae were placed by Schulz (1936) in other tribes. Further studies are needed to determine if other genera. The almost complete lack of ITS (Warwick et al. not paralleled anywhere in the family. simple or no trichomes (only the South African Sisymbrium bruchellii DC. which include Sibaropsis. Koch et al. 2003a) supported the removal of many of its component genera to other tribes. but many South American members evolved branched ones. tetradynamous. 2-lobed stigmas. 2005. Romanschulzieae).104 I. Schulz and Schoenocrambe Greene). this tribe consists of only about 40 species of Sisymbrium (including Lycocarpus O. has branched trichomes). Tribe Sisymbrieae. (1 sp. The Sisymbrieae are characterized by having terete siliques. Indeed. Guillenia and Sibara (Arabideae). As it stands now. The Sisymbrieae was recognized as a tribe of 70 genera and 400 species (Schulz 1936. Koch 2003. 2005a) and ndhF (Beilstein et al. uncoiling of ovate or globose anthers. coiling of linear anthers vs. linifolium) are distributed in Eurasia and Africa. all except one North American species of which (S. Streptanthella. Al-Shehbaz 1988c). The majority of taxa are either glabrous or with simple trichomes. the streptanthoid genera (with crisped or channeled petals. most of Caulanthus.

Thellungiella in the Sisymbrieae subtribe Brayinae.) Rchb. Pachyphragma (DC. incumbent cotyledons.E. ex DC. and Thlaspi s. Schulz (1936) placed Eutrema and Taphrospermum in the Sisymbrieae subtribe Alliariinae. Schulz (2 spp. and a base chromosome number of x=7. 2001b. and yellow flowers.: Systematics and phylogeny of the Brassicaceae 105 chromosome number of x =7. Warwick et al. often pendulous fruits.). 90 species previously assigned to it. auriculate cauline leaves. (9 spp. is unrelated to the ca. Mey. simple or no trichomes. of which the ranges of one species each of Eutrema (10 spp. often palmately veined basal leaves.). Mummenhoff et al.E. (6 spp. entire cauline leaves.str. & Spach. Members of the Thlaspideae are characterized by having simple or no trichomes.I. Sobolewskia M. Bieb. Tribe Isatideae. As discussed above.). and it is likely that the tribe might be unigeneric.) N.) and Myagrum L. 1997b. might belong to this tribe.). The limits of Isatis are controversial. (2003a) suspected that it forms a monophyletic group based strictly on morphology. Al-Shehbaz et al. and the vast majority of those belong to the unrelated Noccaea (ca.) extend into northern North America. Sameraria Desv. Platycraspedum O. and Chalcanthus in the Brassiceae. Tribe Eutremeae. Chartoloma Bunge. Taphrospermum C.) form a monophyletic group (93% bootstrap support) sister to a clade including the tribes Brassiceae. Graellsia Boiss. Platycraspedum in the Lepidieae. Schulz (3 spp. Tribe Noccaeeae. Beilstein et al.). Neomartinella in the Arabideae. (7 spp. As indicated above. This new tribe includes about three genera and some 85 species all except for five New World species of Noccaea (one each in Patagonia and Mexico and three .str. Tribe Thlaspideae. 1.. The tribe consists of 26 species in seven genera restricted to Europe and southwestern Asia. Clearly.A. Members of the Eutremeae are glabrous or with simple trichomes and have white flowers.). and Sisymbrieae.) and Thellungiella O. Peltaria Jacq. O’Kane and Al-Shehbaz (2003) showed the same results regarding the last two genera. Molecular data (Beilstein et al. Busch (3 spp. Thlaspi s. Although not included in any molecular studies thus far and based strictly on the overall morphology. more work is needed in the tribe. In Beilstein et al. & Kir. and a base chromosome number of x=7. & Steud. Tauscheria Fisch. Extensive molecular studies (Koch and Mummenhoff 2001. Chalcanthus Boiss. Both Myagrum and Tauscheria were placed by de Candolle (1821a) with Isatis (including Sameraria) in the Isatideae. Pseudocamelina (Boiss.. (8 spp. it is highly likely that the tribe also includes the genera Pachypterygium Bunge (3 spp.). 85 spp. It is likely that at least some of the genera above would be eventually united with Isatis. it is likely that the total number of species be reduced to a little over 50. Schimpera Hochst. (4 spp. and a base chromosome number of x=7. 1997a. and Thellungiella are nested within Eutrema and are therefore united into one genus (Al-Shehbaz and Warwick 2006b). Spirorrhynchus Kar. (4 spp. (1 sp. 2006) show that Isatis (79 spp. indehiscent. (2 spp. This tribe of over 90 species in eight genera was first recognized by Hayek (1911) as a subtribe of the Arabideae and by Schulz (1936) as a subtribe of the Lepidieae.or 2-seeded. and with critical studies. (1 sp. (2005b) showed that Neomartinella (3 spp.). Parlatoria Boiss. often palmately veined basal leaves. A.).). The differences between the constituent genera of the tribe are minor and rest exclusively on fruit morphology.).. and the monotypic Boreava Jaub. The Isatideae have yellow or rarely white flowers.).). It includes Alliaria (2 spp. Based on differences in fruit compression and cotyledonary position. he included in these suprageneric taxa many unrelated elements. (1 sp.) and immediate allies. 2006) defined the boundaries of this tribe. Koch et al. Glastaria Boiss. This primarily Asian tribe comprises about 25 species. often pinnately divided lower leaves. which has not yet been surveyed. striate or coarsely reticulate seeds. Schizopetaleae. (2006).) and Taphrospermum are sister taxa with 100% support. ex Endl.

A. Koch and Al-Shehbaz (2004). Multicellular stalked glands occur in the Chorisporeae and Anchonieae. (2006). tardily dehiscent siliques. see Gowler. (3 spp. only part of Microthlaspi F. as well as Prantl (1891) and Hayek (1911). (2006). highly supported (97–100% bootstrap) subclades recognized herein as the tribes Hesperideae. Mey. (2006). 1988).). (7 spp. and Diptychocarpus Trautv. Matthiola R. in the Hesperideae and Matthioleae. (Appel and AlShehbaz 2003).Br. Oreoloma Botsch.106 I. The Noccaeeae are characterized by having glabrous. (2 spp. the Hesperideae included many other genera. 1998). Excluding Chorispora R. and the monotypic Zerdana Boiss. Jacquemoud.: Systematics and phylogeny of the Brassicaceae in the United States and Canada) are distributed in Eurasia and northern Africa.K.. Phylogenetic studies of this tribe are in progress (Warwick et al. Although Parrya was not included in Beilstein et al. often auriculate cauline leaves.) should perhaps be united with Sterigmostemum and Clausia. ex DC. de Candolle (1821a. Microstigma Trautv. a genus centered in the Middle East and Europe.Br.. In the study of Beilstein et al. and a base chromosome number of x=7. and it is likely that the absence of such gland occurred independently.). none of them was monophyletic. . Sterigmostemum M. that polytomy included four. and Chorisporeae. Euclidieae. Busch (2 spp. These two tribes were later combined by Janchen (1942) as the Hesperideae and by Al-Shehbaz (1988b) as the Anchonieae. Al-Shehbaz et al.). along with a highly heterogeneous assemblage of others. 34 spp. The Hesperideae includes biennial or perennial plants with simple and/or forked trichomes. and further studies are needed to firmly establish that. 1979) group with Noccaea in a well-supported clade.. The extensive molecular studies [for summaries see Koch (2003). (3 spp. The exact position of Dontostemon and Bunias with the rest of the Anchonieae was not fully resolved in Beilstein et al. Jacquemoud. Tribe Anchonieae. As delimited by Prantl (1891). respectively.. and only a few of his ten segregates might deserve recognition (Koch 2003). However. Bieb. Koch and Bernhardt 2004).) and Pseuoclausia Popov (10 spp. and the remaining segregates should perhaps best treated as synonyms of Noccaea. Hesperis and 20 other genera formed an unresolved polytomy with only 68% support. Koch and Hurka 1999. b) placed Hesperis in the Sisymbrieae..) in North America. with poor representations in northern Africa and central Asia. and Matthiola include species with or without multicellular glands. all of the segregates of Thlaspi by Meyer (1973. (in press) showed that it should be assigned to the same clade including such glands. Hesperis is much in need of thorough systematic and phylogenetic studies. These include Neurotropis (DC. decurrent stigma lobes. smooth seeds. the tribe includes Anchonium DC.Br.K. However. Parrya. Dontostemon.-Trotzky (6 spp.. and Schulz (1936). with only four species of Parrya R. Anchonieae. none of which have uniseriate glands. Schulz (1936) placed all such genera. The tribe is readily distinguished from the rest of the Brassicaceae by having stalked glands with uniseriate stalks terminated with a unicellular gland. By contrast. pers com.).). and the discussion above on Thlaspi] support the clear distinction between Noccaea and Thlaspi. genera of the Anchonieae have multicellular-multiseriate glands. Mey. The Anchonieae include some 12 genera and about 130 species distributed primarily in Eurasia and eastern and northern Africa.). Dontostemon (11 spp. (3 spp.) F.). but in these tribes the stalks are multiseriate and the glands are multicellular. 1984). 1998. Hayek (1911). However. Clausia Korn. (ca. (48 spp. Microthlaspi was subjected to extensive molecular studies (Koch et al. often perennial plants with angustiseptate fruits.. and base chromosome numbers of x=6–10. This unigeneric tribe consists of Hesperis (46 spp. Iskandera N. Tribe Hesperideae. as delimited by all these authors. Yue et al. Bunias L. In addition.

persistent cotyledons larger than the leaves of the plant. but the genus was not included in Beilstein et al.) and Diptychocarpus (1 sp.). though x=9 was reported in some Chorispora. It consists of about 25 genera and over 150 species. Schulz 1936) that the last three genera belong to the Brassiceae. the single species of which. Phylogenetic studies addressing relationships in this tribe are in progress (Warwick et al. Dilophia Thomson (2 spp. but it appeared in a clade of its own in the overall polytomy of the family. The genus formed a monophyletic clade in the work of Mummenhoff et al. though x=8 is less common. (3 spp. well-resolved clades represented by the larger genera Braya and Malcolmia.E. Le Schulz (10 spp.).f. and Vesleskya Opiz.. Sun (1 sp.. members of the Anchonieae also have branched trichomes. often strongly 2-lobed stigmas. The tribe is characterized by the lack of multicellular glands and the presence of incumbent cotyledons.).). and Tetracme Bunge (10 spp. Euclidium R. and their results are taken to support the placement of the genus in its own tribe. Chorispora was placed by de Candolle (1821a. Among these are Diceratella Boiss.). but they need to be studied using molecular data. Atelanthera Hook. has double conduplicate.. Dvora Maresia Pomel (3 spp. entire or 2lobed stigmas. often terete to 4-angled siliques or silicles. branched trichomes (rarely simple or absent)..).g. (1 sp.). (3 spp. (2004a) showed that both Neotorularia and Sisymbriopsis Botsch. but it is generally agreed (e. Leptaleum DC. Shangrilaia Al-Shehbaz. Parolinia Webb (5 spp.. This tribe showed 99% bootstrap support in Beilstein et al.f.). Solms-laubachia Muschl. & Tzvelev (5 spp. ´ onard (11 spp. and a base chromosome number of x=7. ´ k (6 spp. J. erect sepals. Hayek 1911. Malcolmia R. & Hoppe are North American. in the tribe Cakileae. 1821b) with Cakile. erect sepals forming a closed calyx..) Zahlbr. and the monotypic Anastatica L. pers com. Beilstein et al. C. A.Br.).f. Br. circaeoides (L. Al-Shehbaz et al. several genera probably belong to the Euclidieae.. it includes Desideria Pamp. (2006) included only one species of Heliophila.). Warwick et al.). & Kir. multicellular-mutiseriate glands. Yue & H. and a base chromosome number of x=7.). (12 spp. and typically spurred calyx. & Thomson. Morettia DC. This exclusively Asian tribe consists of Chorispora (11 spp. though Matthiola show reductional aneuploid series to x=5 and increase to x=8. Leiospora (C. (2005) with Heliophila s. and Cordylocarpus Desf.A. with 100% bootstrap support. and only seven of 17 species of Braya Sternb. Tribe Chorisporeae. Rapistrum.). moniliform fruits breaking into 1seeded. (2006). Six genera were previously assigned to the Heliophileae (Appel and Al-Shehbaz 1997).l. Notoceras R. Neotorularia Hedge & J.P. and a base chromosome number of x=7. Cryptospora Kar. It is not known if the Heliophileae would remain unigeneric or if it would also include the South African Chamira Thunb.).: Systematics and phylogeny of the Brassicaceae 107 In addition to the multicellular-multiseriate glands. (2005).I. In addition to Braya. (11 spp.Br. This tribe is restricted to South Africa and presently consists of 82 species of Heliophila. and it exhibited internal differentiation into two.). (2006). the New Zealandic Notothlaspi Hook.) might belong here too.) F. (32 spp.). Tribe Heliophileae. but they have recently been united by Al-Shehbaz and Mummenhoff (2005) and Mummenhoff et al. (9 spp. Hayek (1911) and Schulz (1936) placed Chamira and Heliophila . Rhammatophyllum O. (2006).). Cithareloma Bunge (2 spp. Eremobium Boiss. erect sepals. According to Mitchell and Heenan (2000) and Koch (2003). corky segments. It formed a welldefined clade in Beilstein et al. (2 spp. On the basis of morphological grounds. The tribe is primarily Eurasian and northern and eastern African. Tribe Euclidieae.) are polyphyletic. and some of their component species need to be re-assigned to other genera or placed in independent ones. The Chorisporeae is characterized by the lack of branched trichomes and the presence of connivent stigmas.). Mey.

fruit. 2003b).: Systematics and phylogeny of the Brassicaceae in separate tribes. was subjected to extensive molecular studies (Koch 2002. The genus.E. As many as 135 smaller or medium-sized genera. Indeed. 27 spp. simple trichomes or glabrous. A. but he also included genera assigned herein to the Eutremeae (Platycraspedum). Schulz. It is not known if these two genera merit the placement in their own tribe. the systematic position of some of the genera already sampled remain unresolved. entire stigmas. northwestern Africa. and it is distributed primarily in Europe. now Lepidium). All species of the tribe are either glabrous or with simple trichomes. which is distributed primarily in Europe.. . Schulz (1936) recognized the South American tribe Menonvilleae. 1996. (8 spp. Tribe Cochlearieae.) have compound leaves and grow in wet habitats (Koch and Al-Shehbaz 2000). (2006) formed a clade of its own in the overall polytomy of the family. 1999b. as well as five other genera of unknown affinities. now Halimolobos). and trichome morphology. and it too appeared in a clade of its own in the overall basal polytomy of the family.) were placed in the Lepidieae by Schulz. the latter genus fell outside this alliance in the survey of Beilstein et al.) and Cremolobus DC. and six other genera that have not yet been studied molecularly. Other genera.). Al-Shehbaz et al. Although Beilstein et al. Halimolobeae (Poliophyton O. which included Menonvillea DC. Furthermore. but he also included genera now assigned to the Physarieae (Dithyrea) and Aethionemeae (Moriera). Schulz (1936) place Cochlearia in the tribe Lepidieae subtribe Cochleariinae. white petals. Schulz (1936) placed the genus in the tribe Lepidieae subtribe Iberidinae. corymbose infructescences. The genus needs to be studied to check if it belongs to the tribe Smelowskieae. and a base chromosome numbers of x=6 or 7. undivided basal leaves. The Cochlearieae are distinguished by being glabrous plants with rosulate. and the majority have appendaged petals and/or staminal filaments and ‘‘extrafloral nectaries’’ (termed stipules by Marais 1966) at the base of pedicels and/or leaves. Because of the distinctive morphology of Iberis (see below). Tribe Iberideae. including five of Ionopsidium). this tribe consists only of Iberis (ca. and northern North America and Asia. (24 spp. but in the overall phylogeny of the family (Koch 2003). biseriate seeds. 2-seeded fruits. (2006) studied only one species of Ionopsidium. Chromosome numbers are known for only four species of Heliophila. especially in the outer flowers of the corymb. ebracteate racemes. Lepidieae (Stroganowia. and often zygomorphic flowers with strongly unequal pairs of petals. terete or angustiseptate silicles.) and Biscutella (53 spp. the genus is placed herein in a tribe of its own. Both Megacarpaea DC. a feature that has evolved independently in three Australian species of Lepidium (see above). but it is doubtful if they belong to that tribe. remain to be studied for at least one molecular marker.108 I. as do most members of the Cardamineae. with fewer species in northwestern Africa. The Iberideae are distinguished by having strongly angustiseptate. and southwestern and central Asia. the results of Koch and colleagues are taken to support the recognition of this unigeneric tribe. as these taxa show significant similarities in leaf. b) placed them in the Heliophileae. Koch et al. Members of the primarily Chinese Yinshania (13 spp.). representing only about 400 species of Brassicaceae. (2006). The tribe is easily distinguished by having diplecolobal cotyledons. Yinshania appears to be more closely related to the Camelineae than to the Cardamineae. Turkey. and it appeared in a basal polytomy of the family (Koch 2003). but de Candolle (1821a. As presently delimited. (9 spp. This unigeneric tribe consists of Cochlearia (21 spp. but its position with regards to the other groups still needs to be firmly established. often sessile cauline leaves. The single species studied by Beilstein et al. all of which have n=10.

1969. Koch et al. 2002b. and Arabis are taxa with serious problems relating to generic boundaries. hybridized with almost every diploid species with which its range overlapped. Mummenhoff et al. the Brassiceae. 2004). and recent studies involving DNA sequences of microsatellites markers (Sharbel et al. b. which is the most widespread species of the genus. Schizopetaleae. Lo ¨ ve & D. 2002a. a genus distributed primarily in the western United States and Canada. The subject is discussed in further detail in the paper by Marhold and Lihova (2006) in this issue. distinct morphologically and isolated reproductively from both parents. More problematic is the delimitation of taxa the evolutionary history of which involved extensive polyploidy. (2004). Aneuploidy was reported in the B.) strongly suggest that B. Sharbel and Mitchell-Olds 2001). Koch and Al-Shehbaz 2002. As suggested by Dobesˇ et al. However. As indicated by Windham et al. The aim is to apply the knowledge gained from apomictic plants in the potential development of easily propagated apomictic crop plants (Hoisington et al. Pleistocene differentiation in Boechera was greatly influenced by alternating glacial and interglacial cycles. and Lepidium (Lee et al. neither hybridization nor apomixis were addressed. Lo ¨ ve & D.). Widmer and Baltisberger 1999a. 2004) focused primarily on B. Beilstein and Windham 2003. There seems to be evidence that the triploid hybrids also involved three parental species. holboellii (Hornem. 2002. pers. and their ‘‘alleged’’ triploid apomictic hybrid. Roy 1995. the genus appears to be far more complicated than what had been suggested thus far. 2002. Lo ¨ ve. 2001a. 2002a. holboellii species complex (Bo ¨ cher 1951. hybridization. and erroneous determinations in the holdings of the major herbaria can be as high as 40%. Therefore. 2004). 2003c. Boechera consists of over 60 species of sexual diploids (2n=14) and about 40 apomictic triploids (3n=21) (Windham. The pioneering cytological and embryological studies by Bo ¨ cher (1951. and apomixis. Cardamine (Neuffer and Janche 1997. Al-Shehbaz et al. Scheen et al. 1997. and references therein) firmly established apomixis in the genus. . the resulting hybrids often become stabilized apomictic triploids. The molecular basis of apomixis is being investigated in several Boechera species. Preliminary studies (Windham.: Systematics and phylogeny of the Brassicaceae 109 Problematic groups As discussed above. Most of the discussion below will focus on Boechera. van Dijk and van Damme 2000). It is likely that these two phenomena influenced the evolution of all major genera of the family. Franzke and Hurka ´ et al. B. 1983. Franzke and Mummenhoff 1999. Until recently. but molecular studies (summarized in Al-Shehbaz 2003b) show that the two genera are not closely related and placed herein in different tribes. the genus has undergone rapid bursts of reticulate evolution. Grundt et al. 2000. Boechera was considered as part of Arabis (Rollins 1993). Bleeker et al. 2003. ·divaricarpa (A. which produced a mind-boggling array of forms that have blurred species boundaries.I. Mulligan 1996). It appears that in all cases. pers. 1998. 1993. the first serious problem in the study of Boechera is taxon identity. com. Brochmann et al.) A. 2004) suggest that aneuploidy involves non-recombining B chromosomes that may also play a role in apomixis. Sharbel and Mitchell-Olds 2001. A. Sharbel et al. Recent molecular studies (Roy 2001. Lo ¨ ve. stricta (=Arabis drummondii). 2004a. Urbanska et al. 2000. B. Hybridization appears to have played a far more major role in the evolution of the genus than was suggested by Rollins (1983) and Mulligan (1996). Polyploidy and hybridization have been well documented in Draba (Brochmann 1992. and that those are also fixed through apomixis. (2004). Lihova Marhold et al. com. Although Boechera (as Arabis) was subjected to extensive taxonomic and cytological studies (Rollins 1941. 2004 and references therein). 2004b. Franzke et al. 1999. 1992 and references therein. stricta. Nelson) A. Dobesˇ et al.

Beilstein & E. The bibliographical citation and type genera of all tribes are given. nov. basi articulata. chromosome triplication has recently been documented in the entire tribe Brassiceae (Lysak et al. and comparative genomics. Contrary to the listing of Hayek (1911). Kellogg. com. Seven tribes are proposed as new. Herbae suffrutices. perennes vel annuae. chromosome painting. 2005) and useful to a certain extent. The effects of various genes on the development of flower and fruit in the family are covered by John Bowman (2006) and Gu ¨ nter Theissen (2006) in this issue. B. Type genus: Aethionema R. and Al-Shehbaz (1984). (2006). Lepidieae. Brassiceae. 2.) A. in W.Br. and several genera (e. folia integra. Aiton. pers. didymocarpa (Hook. eight tribes (Alysseae. Hortus Kew. Hesperideae.g. much taxonomic emphasis has been placed on the arrangement of flowers in racemes or on solitary pedicels originating from the basal rosette.A. Although genome size is highly variable in the Brassicaceae (Johnston et al.110 I.A. oleracea (n=9) and B. For example. Baum 2002. rapa (n=10) represent ancestral hexaploids. 2005). sessilia vel breviter petiolata. Studies to understand the evolution of the wide array of chromosomal numbers and ploidy levels found in Physaria are in progress (Fuentes-Soriano. ed. Leavenworthia.g. 1812. Mulligan (n=4) and P. Al-Shehbaz et al. it becomes a far more valuable tool when combined with studies involving the determination of chromosome numbers. control the development of rosette instead of raceme flowering in the Brassicaceae (Shu et al.). In fact. Arabideae. Schulz (1936). such as Physaria bellii G. the tribes proposed by de Candolle first appeared in April (de Candolle 1821a) instead of late May (de Candolle 1821b). com. it appears that the entire Brassicaceae evolved after the genome duplication of its ancestors. including LEAFY. What is needed are studies of genome size throughout basal Aethionema to determine if indeed the so-called ‘‘diploid’’ species with n= 7 and 8 are ancient tetraploids. Euclidieae. The recent advancement in genome research and its very promising role in understanding the evolutionary history of the family are covered in the accompanying paper by Lysak and Lexer (2006). Heliophileae. pers. such as B. nigra (n=8). These led to the conclusion that so-called ‘‘diploid’’ species in the genus. As indicated by Kellogg and Bennetzen (2004 and references therein). keys. Selenia) can have both types of flowering on the same plant. to have a similar genome size (Lysak. Taxonomic considerations Of the 49 infrafamilial taxa (19 tribes and 30 subtribes) recognized by Schulz (1936). glabrae saepe glaucae. Yoon and Baum 2004). and complete tribal assignment of all genera of the Brassicaceae are being prepared for a separate publication. T. A synopsis. A. saepe carnosa. Descurainieae) were recognized previously (e. 4: 80. less emphasis should be paid to large-scale surveys such as that of Beilstein et al. We now know that a few genes. Sisymbrieae) are accepted herein.). Fuller understanding of the developmental genetics of various structures will have a major impact in phylogenetic and systematics studies in the Brassicaceae. racemi ebracteati. Far more exciting is comparative genomic studies that focus on evolution within complex groups. This has also been recently suggested by Schranz and Mitchell-Olds (2005). Schulz 1936) as subtribes. The pioneering studies of Lagercrantz and Lydiate (1996) and Lagercrantz (1998) showed that the genomes within the cultivated Brassica underwent extensive triplications of large genomic regions accompanied by chromosomal rearrangements. filamenta .: Systematics and phylogeny of the Brassicaceae Genomics and development There is no doubt that once a clearer picture on the overall phylogeny of the family is obtained. It is rather intriguing to find two species of the same genus. trib. 2000. Tribe Aethionemeae Al-Shehbaz. Gray (n=12). and two (Physarieae.

. Not. basi auriculata vel exauriculata.. Sp. A. Ess. petala alba vel rosea. angustiseptati. glabri. 27: 307. 2: 154. cotyledones accumbentes vel incumbentes. Type genus: Lepidium L. 1975. racemi bracteati vel ebracteati. Type genus: Physaria (Nutt. 2: 650. ovula numerosa. Herbae suffrutices. petiolata. in Ledeb. pilis simplicibus vel multifurcatis.. ´ m. 1847.. Beih. Fl. Tropidocapinae Hayek. Sp. Al-Shehbaz et al. valde compressi. Fam. sessilia vel breviter petiolata. Nat. Tribe Cardamineae Dumort. Tribe Descurainieae Al-Shehbaz. pilis simplicibus vel multifurcatis. folia 1–3pinnatisecta. Tribe Halimolobeae Al-Shehbaz. ´ m. nov. Fl. Syn. nonmucilaginosa. 2: 17. Descurainiinae O. 1: 17. fructus siliquae vel rarissime siliculae. Pl. basi nonauriculata. Tribe Boechereae Al-Shehbaz.A. Tribe Alysseae DC. 1(1): 99. petala flava. Tiefebene: 258. Harmsl. 1893. mucilaginosa. 1821.I.. ovula numerosa. Ital. Nat. Centralbl.. nov.. Schulz in Engler & H. nov. Schizopetaleae subtr. 9: 726. trib. Centralbl. teretes. Char. Tribe Physarieae B. Iles Canaries 3(1): 72. Amer. Kellogg. Fl. fructus siliquae vel siliculae. Mey. 1891.: Erysimeae Dumort. 123. Type genus: Boechera A. Type genus: Arabis L. pilis simplicibus vel furcatis. 1753. Hist. perennes vel annuae. Me Nat. fructus siliquae vel siliculae. Type genus: Descurainia Webb & Berthel. pili furcati praediti vel rarissime glabri. folia integra vel dentata. Sp.. Capselleae Horan. Ital. III. ed. petiolata. ´ m. latiseptati vel teretes. Belg.: 170.. invalid. ovula 18. Tribe Camelineae DC. Lo ¨ ve. ed. Sp. cotyledones incumbentes. Fl. 2. dentata vel integra. . rarissime alba. 1827. racemi saepe ebracteati. Syn. Gen.. Fl. perennes vel annuae. 1830. 1. glabri. Me Nat. Pflanzenfam. Beih. Hist. Herbae perennes vel annuae.L. alba vel purpurea. Syn.A. Austr. teretes. Mus. Syn. cotyledones incumbentes vel accumbentes.: Systematics and phylogeny of the Brassicaceae 111 plerumque alata. mucilaginosa. Herbae perennes vel annuae. racemi ebracteati. folia bi. Type genus: Smelowskia C.. Beilstein & E. Gray. biloculares oligospermi dehiscentes vel uniloculares monospermi indehisentes. Synop. 27: 313. Bot.vel tripinnatisecta. Hist. Pl. latiseptates vel angustiseptates. nov.. Hist. & A. 7(1): 229. semina uniseriata vel biseriata. Stirp. Flora (Regensburg) 19: 410. pilis simplicibus vel multifurcatis. Turriteae Buchenau. folia integra vel rarissime dentata. Me 7(1): 231. 1837. Herbae suffrutices..: Clypeoleae Webb & Berthel. petala alba vel rosea. trib. Hist.. Me Nat. 1848. 1911. N. Pl. 9: 658. Sisymbrieae subtr. Fl. Tribe Arabideae DC. Type genus: Camelina Crantz. Beilstein & E. 1821. Icon. Nat. cotyledones incumbentes. 2: 643. Mus. nonmucilaginosa. ovula numerosa. 1936.. Illustr. 1762. Fl. Mus. Nat. Gray.A. Type genus: Halimolobos Tausch. Hist. ovula 2 vel numerosa. 1753. semina uniseriata vel biseriata. Kellogg. Pl. Nat. A. basi saepe auriculata. 2: 654. 7(1): 239. Iles Canaries 3(2. 1: 162. basi nonauriculata..1): 89. 1753. fructus siliculae. 1827. Lo ¨ ve & D. 2: 664.E. trib. 1821 (as Lepidineae).: Lepidieae subtr. ´ m. semina uniseriata vel biseriata. Tribe Lepidieae DC. Beilstein & E. Type genus Alyssum L. 1836. 128: 513. 1895. 1: 100.: Schizopetaleae subtr. Mus. 1976. Amer. Nordwestdeut.. Syn. 1753. 1893. Pflanzenfam. semina uniseriata vel biseriata.: Selenieae Torr. Type genus Cardamine L. Physariinae Prantl in Engler & Prantl. fructus siliquae vel siliculae. nom.. Belg. angustiseptati vel teretes. Beilstein & E. N. Kellogg. racemi ebracteati vel bracteati. 1821. Lyrocarpinae Hayek. Nasturtieae Caruel in Parl. petala flava. 7(1): 240. publ. Bot.. Kellogg.A. trib.) A.: Cardarieae Caruel in Parl. Tribe Smelowskieae Al-Shehbaz. 1894. Pl. Syn. Bot. 1911. 1836. sessilia. Rob.: 124. 1838. 17B: 649.

pilis simplicibus.Br. ed. 2: 657. fructus siliquae vel siliculae.. Altaic. Me ´ m. 7(1): 242.: Peltarieae Caruel in Parl. Tribe Cochlearieae Buchenau. Tribe Iberideae Webb & Berthel. nonmucilaginosa. Type genus Heliophila L.. saepe palmativenosa.. 244. Sp. ´ m.. Nat. Schulz in Engler & Harms. A. saepe auriculata. ed. ´ m. 1894. Sp. Chlor. quadrangulares vel compressi. Mus.. 2.. Hist. 1895. 1821. 1821. glabrae vel rarissime pilosae. ´ m. Me ´ m. Synop. (Paris) 7(1): 237. Pflanzenfam. Herbae perennes vel annuae. Nat. 9: 1043. Me Hist. Ital. Bot. Hist... Centralbl. Centralbl. trib. 2: 648. (Paris) 7(1): 247. Sp. Me 7(1): 236. 2. 2: 647. ovula 4 vel numerousa. Mey. Bot. pilis simplicibus. Pflanzenfam. Aiton. trib. Matthioleae O. Hist. Me Nat. Fl. 1845. semina uniseriata.: Thelypodieae Prantl in Engler & Prantl... Type genus: Anchonium ´ m. 9: 1029. 17B: 557.E. 1821. 1821. Type genus: Noccaea Moench.: Lepidieae subtr. laevia. nom.: Myagreae Caruel in Parl. 1891. Hist. Type genus: Iberis L. 7(1): 236. Mus. 2: 666. Pl. 1821. 1821.: Systematics and phylogeny of the Brassicaceae ´ m. 1821. invalid. Pl.. Sp. glabrae vel rarissime pilosae.. auriculata vel nonauriculata. 7(1): 242. Chorispora R. Syn. DC.. Sp. Nat. 1753.A. ed. Beilstein & E.. Beih. 1911. nov. 1891. Hortus Kew. 2: 154. ´ m. 2: 670. Syn. 2. Me Nat. Mus. 27: 315. fructus siliculae. ed. nom. Syn. Nat.. 17B: 298.. Type genus: ´ m. valde compressi. 7(1): 244. James Beck. 1821. Type genus: Hesperis L. Type genus: Schizopetalon Sims. Mus. 27: 315. 3: 104. semina uniseriata. 7(1): 237. Me Nat. 89. Syn. 2. ´ m. Type genus: Isatis L. glabri. Nat. 1936. Tribe Isatideae DC. Tribe Anchonieae DC. Iberidinae Hayek. 4: 74. petala alba vel rosea. 1821. Nat. sessilia. 2379. Tribe Schizopetaleae R. Robins. Mus.E. Al-Shehbaz et al. Erucarieae DC. Mus. 1893.. 2: 163. Sara Fuentes- . Tribe Noccaeeae Al-Shehbaz. 1911. Pflanzenfam.A. 2: 645.. 1821. Hist. 1823.: Sinapeae subtr. The authors are grateful to Oliver Appel. Nat. & Bunge. Raphaneae DC. III. neae DC. Hist. Syn. Hist. Me Nat. Stanleyeae B. Hist. Mus.. Nat. Mus. Nat. Nat.T. Nat. in Barne Ann. 7(1): 243.1): 92. Mag. Me 1821. Nat. Me ´ m. nom. Me 7(1): 241. III. Hist. Mus. 17B: 300.Br. Mus. Nordwestdeut. Nat. invalid. Ital. 1753. C. 1821. 16: 68. 2: 155. Pringleeae Hayek.. Me Nat. Nat. Syn. Type genus: Thlaspi L. Mus. 1: 105. ´ m. ´ oud. 1821.. Nat.: Brachycarpaeae DC.. Herbae perennes vel annuae. III. 2: 926. folia integra. Fl. ´ m. 7(1): 246. nov. in W. Syn. Suppl. 7(1): 234.112 I. ex DC. 1821(as Cakilineae).Br. Hist. Mus. Kellogg. Fl. ed. folia integra. Hist. Tribe Thlaspideae DC. 1831. Tribe Chorisporeae Ledeb. Iles Canaries 3(2... Hist. Pflanzenfam. Amer.L. Pflanzenfam. Type genus: Euclidium R. Melvill. Meth. N. Tribe Sisymbrieae DC. 1891. 1823. Bot. invalid. Cochleariinae Prantl in Engler & Prantl.: Buniadeae DC. Nat. Tribe Hesperideae Prantl in Engler & Prantl. 1821. Me 7(1): 236. Pl. Hist. 1753. 1763. Schulz in Engler & Harms. Hist. Nat. Mus. 1893. ´ m.: Anastaticeae DC. basalia longe petiolata. 1753.. 1812. Romanschulzieae O. Sp. Kellogg. Hist. Me Mus. C. 1802. Me Nat. Type genus: Sisymbrium L. 7(1): 1821. teretes. 1821. 9. Mus. cotyledones incumbentes. angustiseptati... Schulz in Engler & Harms.. Syn.. Pl. ´ m.A. PsychiMe ´ m. Hist. 1936... Zilleae DC. Me Nat.. Pflanzenfam. Hist. Mus.. 2: 663.E. Type genus: Brassica L. Sp. Pl. Tribe Euclidieae DC. 1837. Mus. Nat.. Velleae DC. folia caulina sessilia vel petiolata. Tiefebene: 245. Tribe Eutremeae Al-Shehbaz. (Paris) 7(1): 242. Pl. glabri.. Mus. Me Nat. Beilstein & E. t. Donovan Bailey. Beih.. petala alba vel rosea. Tribe Brassiceae DC. Hist. 1753. Type genus: Eutrema R.: Cakileae DC. 1753. 1753.Br. Pl.. Mag. Fl. Streptantheae O. 1936. 7(1): 245. Fl. Tribe Heliophileae DC. 1821. Pl. ´ m. 7(1): 246. ´ m. Sp. Type genus: Cochlearia L.. Syn. 7(1): 245. Hist.

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