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Five Additional Tribes (Aphragmeae, Biscutelleae, Calepineae,
Conringieae, and Erysimeae) in the Brassicaceae (Cruciferae)
Author(s): Dmitry A. German and Ihsan A. Al-Shehbaz
Source: Harvard Papers in Botany, 13(1):165-170. 2008.
Published By: Harvard University Herbaria
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Extensive molecular studies (for leads, see
Koch et al., 2003; Mitchell-Olds et al., 2005;
Al-Shehbaz et al., 2006; Beilstein et al., 2006;
Bailey et al., 2006; Koch et al., 2007; and
Warwick et al., 2007) were conducted during the
past dozen years on the phylogeny and evolu-
tion of Brassicaceae (Cruciferae). As a result,
tribal classifications of the family based strictly
on morphology, such as the systems of de
Candolle (1821), Prantl (1891), Hayek (1911),
Schulz (1936), and Dorofeyev (2004), as well
as the modifications of Schulz’s system by
Janchen (1942) and Al-Shehbaz (1984), were
shown to be highly artificial. Although the
boundaries of only a few tribes (e.g., Brassiceae
DC., Heliophileae DC.) remained largely unal-
tered, circumscription of almost all of the
remaining tribes needed re-adjustments.
The first attempt to achieve a tribal classifi-
cation of the family based on molecular studies
and re-examination of morphology was made by
Al-Shehbaz et al. (2006). In that study, 25 tribes
(including six new) were recognized. However,
further adjustments at the tribal level, including
placements of several smaller genera not previ-
ously studied or with unresolved positions on
the phylogenetic trees (see Bailey et al., 2006;
Warwick et al., 2007) or super network (Koch
et al., 2007), became immediately evident. As a
result, the tribes Dontostemoneae Al-Shehbaz &
Warwick, and Malcomieae Al-Shehbaz &
Warwick were described as new, and the tribe
Buniadeae DC. was re-established (Al-Shehbaz
and Warwick, 2007).
Based on molecular studies by German et al.
(in prep.), the present paper deals with the tribal
assignments of nine, predominantly Old World,
genera (Aphragmus Andrz. ex DC., Biscutella
L., Calepina Adans., Conringia Heist. ex Fabr.,
Erysimum L., Goldbachia DC., Megadenia
Maxim., Spirorrhynchus Kar. & Kir., and Zuvanda
(Dvořák) R. K. Askerova), either inadequately
studied or with no previous molecular studies.
They are placed in five tribes, of which two
(Aphragmeae and Conringieae) are described as
new, and three (Biscutelleae, Calepineae, and
Erysimeae) are re-established.
FIVE ADDITIONAL TRIBES (APHRAGMEAE, BISCUTELLEAE,
CALEPINEAE, CONRINGIEAE, AND ERYSIMEAE)
IN THE BRASSICACEAE (CRUCIFERAE)
DMITRY A. GERMAN1AND IHSAN A. AL-SHEHBAZ2
Abstract. The tribes Aphragmeae and Conringieae are proposed as new, and the tribes Biscutelleae, Calepineae,
and Erysimeae (Brassicaceae; Cruciferae) are re-established. The unigeneric Aphragmeae, Biscutelleae, and
Erysimeae include the genera Aphragmus, Biscutella, and Erysimum, respectively. The Conringieae comprises
Conringia and Zuvanda, whereas the Calepineae includes Calepina, Goldbachia, and Spirorrhynchus. The prior
tribal assignments of these genera, total number of species, and geographic distribution of all five tribes are listed.
Keywords: Brassicaceae, Cruciferae, tribal classification
We are grateful to GustavoA. Romero-González and Donna Tremonte for their editorial advice and help. We thank Steve
L. O’Kane, Jr. and an anonymous reviewer for their remarks on the manuscript.
1South-Siberian Botanical Garden, Altai State University, 656049, Lenin str. 61, Barnaul, Russia. Email: oreoloma@
2Missouri Botanical Garden, 2345 Tower Grove, St. Louis, Missouri 63110, U.S.A. Email: firstname.lastname@example.org.
Author for correspondence.
Harvard Papers in Botany, Vol. 13, No. 1, 2008, pp. 165–170.
© President and Fellows of Harvard College, 2008.
This genus previously included about four
species (Ebel, 1998), but has recently been
expanded by the discovery of new species and
by its union with the Himalayan Staintoniella H.
Hara and Lignariella Baehni (Al-Shehbaz,
2000, 2002, 2003; Warwick et al., 2006a). As
presently delimited (Warwick et al., 2006b), the
genus includes 11 species, centered primarily in
the Himalayas and adjacent central Asia, with
one species, Aphragmus eschscholtzianus Andrz.
ex DC., distributed in the northern Russian Far
East,northwestern Canada, and Alaska.
Aphragmus was placed in the tribes
Arabideae DC. and Sisymbrieae DC. by Hayek
(1911) and Schulz (1936), respectively. It was
not assigned to a tribe by Al-Shehbaz et al.
(2006) because none of its species were
included in previous molecular studies.
Although Warwick et al. (2006a) clarified the
generic limits of Aphragmus (8 spp.) and
showed that it is unrelated to Braya Sternb. &
Hoppe (tribe Euclidieae DC.), they did not
address its tribal position. Warwick et al.
(2006a) showed that Aphragmus formed a dis-
tinct, well-supported clade separate from the
other tribal groups (Arabideae, Eutremeae, and
Euclideae) included in the analysis. Koch et al.
(2007) included only A. oxycarpus (Hook.f. &
Thomson) Jafri and showed it to form a weakly
supported clade with a species of Heliophila L.
in a basal polytomy of the family-wide tree.
That relationship was not confirmed in the ITS-
based phylogeny (German et al., in prep.), in
which both A. eschscholtzianus and A. involu-
cratus (Bunge) O. E. Schulz formed a well-sup-
ported clade (100% bootstrap [BS]) distinct
from the other existing tribes.
Aphragmus is characterized by an indumen-
tum of minute simple and forked trichomes,
entire or 3–7-lobed leaves, bracteate inflores-
cences, usually eseptate fruits, and often
minutely papillate seeds on long filiform funi-
cles often longer than seeds (except for four
species previously assigned to Lignariella). This
character combination is not found in other gen-
era of the family and, together with its molecu-
lar distinctness (German et al., in prep.), support
the placement of Aphragmus in an independent,
unigeneric tribe, Aphragmeae.
166 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
BISCUTELLA AND MEGADENIA
Biscutella (53 spp.; Warwick et al., 2006b), is
a primarily Mediterranean genus placed by de
Candolle (1821) in the tribe Thlaspideae DC.,
and by Hayek (1911) and Schulz (1936) in the
tribe Lepidieae DC. Such placements were
based primarily on having angustiseptate silicles
characteristic of all three taxa. Molecular phy-
logenetic studies by Beilstein et al. (2006)
included only one species of Biscutella, whose
position was not resolved in the tree and, as a
result, the genus was not assigned to any tribe
by Al-Shehbaz et al. (2006).
The monospecific genus Megadenia includes
M. pygmaea Maxim., a species distributed in S.
Siberia, S. Russian Far East, and C. China (E.
Xizang, S. Gansu, Qinghai, and W. Sichuan).
The first molecular study of the genus by
German et al. (in prep.) demonstrated that it
forms a moderately supported clade (71% BS)
with Biscutella. Both genera have didymous sil-
icles that split at maturity into two one-seeded
mericarps. Although this fruit type has evolved
independently in Lepidium L. (Lepidieae),
Heliophila L. (Heliophileae DC.), Megacarpaea
DC. and Cremolobus DC. (both unassigned to
tribes), Physaria (Physarieae), and Physoptychis
(Alysseae), the clade containing Biscutella and
Megadenia was not related to any of them.
Therefore, Biscutella and Megadenia are
assigned herein in the re-established tribe
Biscutelleae Dumort. Dorofeyev (2004) recently
placed all genera of the family with didymous
fruits in the tribe Cremolobeae. In doing this, he
ignored all the morphological, cytological, paly-
nological, and molecular data that clearly sup-
port the independent origin of didymous fruits
at least five times in the family (Al-Shehbaz et
CONRINGIA AND ZUVANDA
Conringia, a Eurasian genus of six species
centered primarily in Turkey and adjacent Iran,
was placed by Hayek (1911), Schulz (1923,
1936), and Al-Shehbaz (1985) in the tribe
Brassiceae. Its tribal assignment was based pri-
marily on having in one species, C. planisiliqua
Fisch. & C. A. Mey., subconduplicate cotyle-
dons, a feature characteristic of most members
of the Brassiceae. The tribe was subjected to a
number of phylogenetic studies (see Warwick
and Sauder, 2005, and references therein), but
the placement of Conringia remained unresolved
after its exclusion from the Brassiceae (Lysak et
al., 2005; Al-Shehbaz et al., 2006). Conringia
showed strong alliance (92 to 100% BS) to the
tribe Noccaeeae Al-Shehbaz, Beilstein & E. A.
Kellogg in the ndhF (Beilstein et al., 2006) and
ITS studies (Bailey et al., 2006; German et al.,
in prep.), and similar support (90% BS) in the
trnL-F phylogeny of Koch et al. (2007).
The southwest Asian Zuvanda (3 spp.) was
segregated from Malcolmia W. T. Aiton by
Askerova (1985) based on its lack of branched
trichomes and presence of auriculate leaves.
Malcolmia was placed by de Candolle (1821),
Hayek (1911), and Schulz (1936) in the tribes
Sisymbrieae, Alysseae DC., and Hesperideae
DC., respectively. More recently, Dorofeyev
(2002) transferred all species of Zuvanda and
Conringia perfoliata (C. A. Mey.) N. Busch to
Moricandia DC., but that transfer was not fol-
lowed by Appel and Al-Shehbaz (2003) and Al-
Shehbaz et al. (2006) because Moricandia, like
other members of Brassiceae, has conduplicate
cotyledons (vs. incumbent cotyledons in
Zuvanda). As shown by Lysak et al. (2005) and
Warwick and Sauder (2005), Moricandia is a
member of Brassiceae unrelated to Conringia.
Warwick et al. (2007) demonstrated that
Zuvanda crenulata (DC.) R. K. Askerova, Z.
exacoides (DC.) R. K. Askerova, and
Goldbachia laevigata (M. Bieb.) DC., formed a
separate, but loosely supported (< 50% BS)
clade, unrelated to Malcolmia, and the other 13
tribes included in their study; but they did not
assign the three species to a tribe. Although they
did not include any member of the Brassiceae in
their study, Zuvanda was unrelated to the
Sisymbrieae, a tribe most closely related to the
Brassiceae (Al-Shehbaz et al., 2006). German et
al. (in prep.) showed that Conringia perfoliata
and Z. crenulata formed a clade related to, but
distinct from, the Noccaeeae.
Both Conringia and Zuvanda resemble mem-
bers of the Noccaeeae in being glaucous and
glabrous or with only simple trichomes, and in
having entire or dentate, sessile, mostly amplex-
icaul, cauline leaves. However, they differ by
having terete, latiseptate, or 4-angled (vs. angus-
tiseptate) and linear siliques (vs. oblong, elliptic,
obcordate, obovate, or obdeltoid silicles), many
(vs. few) seeds per fruit, and often decurrent (vs.
entire) stigmas. These significant differences
justify the placement of Conringia and Zuvanda
in a tribe separate from the Noccaeeae and rec-
ognized herein as Conringieae.
2008 GERMAN AND AL-SHEHBAZ, BRASSICACEAE (CRUCIFERAE) 167
Calepina, a monospecific southwest Asian
genus, was placed by de Candolle (1821) in the
tribe Zilleae and by Hayek (1911), Schulz
(1919, 1936), and Al-Shehbaz (1984) in the tribe
Brassiceae because of having slightly condupli-
cate cotyledons with involute margins. Gómez-
Campo (1980) suggested the removal of
Calepina from the Brassiceae, but he later
(Gómez-Campo, 1999) retained it in that tribe.
Recent molecular studies (Lysak et al., 2005;
Warwick and Sauder, 2005) clearly supported
the exclusion of Calepina from the Brassiceae,
a position followed byAl-Shehbaz et al. (2006),
but no tribal placement was suggested.
Goldbachia, a Eurasian genus of six species,
was placed by de Candolle (1821) in the tribe
Anchonieae DC.; by Hayek (1911) in the tribe
Arabideae; and by Schulz (1936) in the tribe
Hesperideae. Recent molecular studies by
Beilstein et al. (2006) revealed a weak relation-
ship between Goldbachia and the tribe
Thlaspideae. Both Goldbachia and Zuvanda
showed no affinity to any of the 13 tribes
included in the analysis (Warwick et al., 2007).
Goldbachia was not assigned to a tribe by
Al-Shehbaz et al. (2006).
The Asian monospecific Spirorrhynchus was
placed by Hayek (1911) in the tribe Arabideae
and by Schulz (1936) in the tribe Euclidieae.
The genus was not included in previous molec-
ular studies, though Al-Shehbaz et al. (2006)
suspected affinity to the tribe Isatideae.
Molecular studies by German et al. (in prep.)
showed that Calepina,Goldbachia, and
Spirorrhynchus formed a clade (58–67% BS)
not closely related to any of the tribes. They are
placed herein in the re-established Calepineae
Horan., a tribe characterized by several charac-
ters given below.
Calepina differs from both Goldbachia and
Spirorrhynchus in having spreading, non-sac-
cate sepals, somewhat zygomorphic petals,
basally non-articulate pedicels, and subcondu-
plicate cotyledons (Appel and Al-Shehbaz,
2003). The three genera, however, resemble
each other in habit, indumentum, cauline leaves,
and fruits (see below).
Erysimum comprises some 223 species
(Warwick et al., in prep.) distributed primarily
in Eurasia, with 8 species native to NorthAfrica
and Macaronesia, and 16 to North America
(authors’ compilation). Many authors recog-
nized Cheiranthus L. and Erysimum as distinct
genera distinguished by the presence of median
nectaries (vs. their absence), terete or 4-angled
(vs. latiseptate) fruits, and incumbent (vs.
accumbent) cotyledons. These differences were
considered by some to be significant at the tribal
level. Indeed, de Candolle (1821) placed
Erysimum and Cheiranthus in the tribe
Sisymbrieae and Arabideae, respectively,
whereas Hayek (1911) placed Erysimum in the
Arabideae and Cheiranthus in the Alysseae;
Schulz (1936) maintained both genera in the
Al-Shehbaz et al. (2006) assigned Erysimum
(including Cheiranthus) to the broadly circum-
scribed tribe Camelineae DC., a position that
was later maintained by Warwick et al. (2007).
The genus Erysimum formed a distinct mono-
phyletic group in several ITS-based studies
(e.g., Heenan et al., 2002; Warwick et al. 2007).
Molecular studies by German et al. (in prep.)
demonstrated that the Camelineae sensu Al-
Shehbaz et al. (2006) is polyphyletic, and
Erysimum (3 spp.) formed a separate, well-
resolved clade from other members of the
Camelineae, that merits the recognition at the
tribal level, herein Erysimeae Dumort. In their
comprehensive ITS phylogeny of the
Brassicaceae, Bailey et al. (2006) also showed
Erysimum to form a well-resolved clade with
100% bootstrap support.
168 HARVARD PAPERS IN BOTANY Vol. 13, No. 1
Based on the above discussions, the follow-
ing five tribes are recognized. The number of
species in a given genus follows Warwick et al.
(2006b). Listing of the genera under each tribe
is based on molecular studies that are either pub-
lished or in preparation.
Tribe Aphragmeae D. German & Al-Shehbaz,
trib. nov. Type genus: Aphragmus Andrz. ex
DC., Prodr. 1: 209. 1824.
Herbae perennes vel annuae, pilis minutis
simplicibus vel furcatis; folia basalia integra vel
3–7-partita; folia caulina petiolata vel sessilia,
basi cuneata nonauriculata; racemi bracteati;
petala alba, rosea, azurea vel purpurea; ovula
numerosa; fructus siliculae vel siliquae, glabri
vel pubescentes, tereti vel latiseptati; stigma
integra; semina uni- vel biseriata, rarius soli-
taria; cotyledones incumbentes.
Annual or perennial herbs; trichomes minute,
simple or forked; basal leaves entire or
palmately 3–7-lobed; cauline leaves petiolate or
sessile, cuneate, non-auriculate at base; racemes
bracteate; petals white, rose, deep-blue, or pur-
ple; ovules numerous; fruits siliques or silicles,
glabrous or pubescent, terete or latiseptate;
stigma entire; seeds uni- or biseriate, rarely soli-
tary; cotyledons incumbent.
This unigeneric tribe includes 11 species dis-
tributed primarily in the high mountains of cen-
tral Asia and Himalaya, and only Aphragmus
eschscholtzianus grows in the arctic (see above).
The tribe is distinguished from other tribes by a
combination of bracteate racemes, minute tri-
chomes, numerous ovules, entire stigmas, and
Tribe Biscutelleae Dumort., Fl. Belg.: 118.
1827. Type genus: Biscutella L., Sp. Pl. 2: 653.
Annual or perennial herbs; trichomes simple
or absent; basal leaves entire, dentate, or lobed;
cauline leaves sessile, often non-auriculate at
base, sometimes absent; racemes bracteate or
not; petals white or yellow; ovules 2; fruits sili-
cles, didymous, glabrous or pubescent, angus-
tiseptate; stigma entire; seeds flattened;
A bigeneric tribe consisting of the primarily
Mediterranean Biscutella (53 spp.) and the
monospecific Megadenia. Schulz (1936) placed
both genera in the tribe Lepidieae subtr.
Iberidineae, but this subtribe included a hetero-
geneous assemblage of genera assigned by Al-
Shehbaz et al. (2006) to at least four tribes. It is
distinguished from other tribes by the didymous,
angustiseptate silicles, 2-ovuled ovaries, entire
stigmas, and accumbent cotyledons.
Tribe Calepineae Horan., Char. Ess. Fam.: 169.
1847. Type genus: Calepina Adans., Fam. 2:
Annual or biennial herbs; trichomes simple
or absent; basal leaves rosulate or not, dentate or
pinnatifid to pinnatisect; cauline leaves petio-
late or sessile and auriculate to amplexicaul;
racemes bracteate or not; petals white, pink,
purple, or yellowish; ovules 1–3; fruits silicles
or siliques, indehiscent, woody, often glabrous,
terete, or slightly flattened; style short or beak-
like; stigma entire; seeds aseriate; cotyledons
incumbent, rarely subconduplicate.
The tribe consists of eight Asian species, of
which six belong to Goldbachia and one each to
Calepina and Spirorrhynchus. It is distinguished
from other tribes by the indehiscent, woody
fruits, 1–3-ovuled ovaries, simple or no tri-
chomes, and entire stigmas.
Tribe Conringieae D. German & Al-Shehbaz,
trib. nov. Type genus: Conringia Heist. ex Fabr.,
Enum.: 160. 1759.
Herbae annuae vel biennes, pilis simplicibus
vel nullis; folia basalia integra vel dentata; folia
caulina integra vel dentata, sessilia, basi cor-
data, sagittata vel auriculata; racemi
ebracteati; petala alba, lutea, rosea, vel pur-
purea; ovula numerousa; fructus siliquae,
glabri, tereti, latiseptati, vel angulari; stigma
capitata vel conica, biloba, decurrentis, raris-
simi integra; semina uniseriata, cotyledones
incumbentes vel rarissimi subconduplicati.
Annual or biennial herbs; trichomes simple
or absent; basal leaves rosulate or not, entire or
dentate; cauline leaves entire or dentate, sessile,
cordate or sagittate to auriculate at base;
racemes ebracteate; petals white, yellow, rose,
or purple; ovules numerous; fruits siliques,
glabrous, terete, latiseptate, or angled; stigma
capitate or conical, 2-lobed, decurrent, rarely
entire; seeds uniseriate; cotyledons incumbent
or rarely subconduplicate.
The tribe includes Conringia (6 spp.) and
Zuvanda (3 spp.).All species grow in southwest
Asia, but the range of one, C. planisiliqua,
extends into the Himalayas, whereas C. orien-
talis (L.) Dumort. is a naturalized weed outside
its native Eurasian range. The tribe Conringieae
is distinguished from other tribes by being pri-
marily glabrous or with simple trichomes only
and by having sessile, cordate, auriculate or
sagittate cauline leaves, linear fruits, capitate or
conical and decurrent stigmas, and incumbent
or rarely subconduplicate cotyledons.
Tribe Erysimeae Dumort., Fl. Belg. 123. 1827.
Type genus: Erysimum L., Sp. Pl. 2: 660. 1753.
Annual or perennial herbs, sometimes shrubs
or subshrubs; trichomes sessile, malpighiaceous
or stellate; basal leaves rosulate, entire or den-
tate, rarely pinnately lobed; cauline leaves entire
or dentate, petiolate or sessile, non-auriculate or
very rarely auriculate at base; racemes often
ebracteate; petals white, yellow, rose, or purple;
ovules numerous; fruits siliques, often pubes-
cent, terete, latiseptate, or angled; stigma capi-
tate, 2-lobed or subentire; seeds uniseriate or
rarely biseriate; cotyledons incumbent or
This unigeneric tribe consists of 223 species
in Erysimum. It is distinguished from the other
tribes by the predominantly malpighiaceous and
stellate trichomes with unbranched rays, many-
seeded siliques, and predominantly yellow flow-
ers. Both Erysimum and Cheiranthus were
simultaneously published by Linnaeus (Sp. Pl.
2: 660, 661. 1753). Wettstein (1889) was the
first to unite the two genera under Erysimum
and, therefore, this generic name has priority
(McNeill et al., 2006: Article 11.5).
2008 GERMAN AND AL-SHEHBAZ, BRASSICACEAE (CRUCIFERAE) 169
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