Table Of Content*
Volume 90
Annals
Number
3
of the
2003
Missouri
Botanical
Garden
SYSTEMATICS OF EURASIAN
AND NORTH
AFRICAN
DORONICUM
(ASTERACEAE:
SENECIONEAE)'
Doionicum e<.mpii*e* herbs distributed Kurope. Nor
Tin- -.-nil- \*lera< r;ic: Si-iii-iiniK ill m ia in
i < 1 1 1 1 I
I |
A
Asia. worldwide revision ol the genus recognizing 2(> specie* and subspecies is presented. In llie pies
I
treatment no age m .gm/ed. Sewn name* .m- newh lectotypified herein: Arnica dor
nl
i
i i i
Doronicum caucasicum M. Biek. Doronicum politic l.halierl. Doroiucurn scorpioides Lam.. Doromcuin
s
Dominium
The genus Asteraeeae: Seneci- This genus belongs the Senecioneac, one of
in
1..
1
complex
oneae) includes rhizomatous herbs with vellow or the largest and most tribe* in the Astera-
green-tinted radiate capitula. All phvllaries are eeae with 2T genera and around .'.200 specie*
1
&
Bentham
similar, general herbaceous ami arranged in Inn (Cassini, 1819; Hooker. I»7:il>: Hoff-
l\
or three rows. Cvpselac are cviindrie to obowile- mann. 1892: Nordenstam. 1977; Bremer. 1994). Its
20 presmnablv group
c\ linen, nidi 10 :»mj liidnia rihs anil bear a pap- species constitute a ualural
mimileh scabrous capdlan (Bremer. 1994; involucre without shorter supple-
of wliite-linled
[)iis
bristles. Tin- ran be absent in ray flowers mi-nlaiA bracts, phvllaries herbaceous arranged in
|>,i|)|ius
of some heterocarpic species. two or three rows, and cspselac cviindrie to ob-
the manuscript, the curators of AV, B. B-W. BC. BCK BM. BOLO. BR
1
OKM. OZL. H\K. IKAY JACA,
BOIS. (;-I)C. GAZI. (iK. Oil. JK. K. ,\l.
MAF, MO. NAP. NY. P-HA. IMA. BO. S. SAM'. S/.B. UPS. W, WU, ZA
.1.— ii-
I
Garmendia for their help with nomencla-
ml matters, a* well a* Juan (.aslillo lor the d ,„-.
.1
supported by grant 4.S 'B'W,-(
)( l.'i l'«; .,1
I I
Sj)a-ii*li Hieceiou Oeneial de Knsenan/.a S e Inwsligaeion Oientffica.
2 Real Jardfn Botanico. CS1C. Mini •:-2B(H Madrid. Spain. ines(«>ma-rjb.csie.os
I'la/a d,-
I
Ann. Missouri Bot. Card. 319-389. 2003.
90:
presented \ppendiccs and
e in 2. respectiv el
I
made
Observations were
rica lo Asii directly or with the a
rocky place
8
from sea level lo 5000 in of elevation. in and cvpselae were studied by SKM. Quant
Although there are several regional studies of the lative characters were recorded u-iug a Brown
genus (Turkey, Kdmondson, 1973, 1975, 1978; Ar- Shape- Plus digital caliper (model 599-571-3). Me,
c.
made
/........!< U(Tr on herbarium specimens,
menia. Avetisvan. ( >i!0; Ihciiau peninsula. ha- a
<
I
and
con. 1987: Belgium. Duv igneaud. 1992). there r flattening drying.
is
maps
each
Distribution
only one previous worldwide revision of Doronicum for
Note geogiaph
new idied. that
(Cavillier. 1907. 1911). Fifteen speeies (l)iels.
1022: Widder. 1925: Sergievskaja. 1949; Widder
&
Fdmondson.
Bechinger. 1950: 1973. 1978: Cha-
&
con, 1987; Perez I'enas. 1990; Perez 1994;
et al..
&
Chen. 1998) and hybrids (Bornmuller Koch.
six
1030; Widder. 1031, 1918; Stare. 1991) were sub-
Two
sequently described. species inehided Cav-
in
monograph have since hern transferred
illier's to
the Arabic word "darawnay." used two
for least
al
other genera hool.cri C. Clarke e\ look.
(i.e.. B.
I),
I
The
different plants (Dozy. 1877). pre-Finnaeai.
(Kitamura. and
to :\<innoglollis 1980). D. thibctan-
botanists (Dioscorides. 1551. 1557; Dodoens. 1571)
um &
Cavill. lo Aster Alvarez Fernandez Nielo Fel-
(
Do
and (week
other authors referred species
to of
iner. 2(KM))). Cavillier (1907. 1911) studied the
ronicum as \conitum pardal'mm ami plant
lies, llie
morphology genu-
of ureal detail. especially
in
1 1 1 .
was probably ml roduced Western culture Av-
in by
the uidumenlum. Ins proposed infrageneric
!>ul
irenna (Dodoens. 1574: Dalenrhamps,
1587).
classification of rather limited value since these
is
The genus Doronicum was described by Lin-
groups are ». uielv defined based mainly on non-
<>l
naeus (1753) to include four species, only two of
which arc currently accepted the genus: D. par-
in
any infragenerir framework
in is
and The remaining
D. plantagineuin.
i/iilioiiclirs
species correspond Scnccio and respec-
to \stcr.
tively. Further, one species of \rnicd described bv
l.inuaeiis 753i. scorpioides also belongs
in
I. I...
( I
The objectives were
rk.
Doronicum
as recognized by l.amarck (1780). Sev-
lorphological characters
(quali-
eral pre-Finnaean authors also confused species
possible so that (I) only
Donmicum
of Scnccio. Aster, and Arnica wild
(Dalenehamps. 1587: Clusius. 1601: Bauhin.
1023; Tournefort. 1700). In particular, the overall
morphological between \mi<n and Do
similarity
ronicum suggested their close affiliation until the
1970s. However. Nordenslanfs (1977) micromor-
phological study of style, anthers, and pollen de-
finitively has excluded \rnicu from the Seneci-
More than 50 qualitative and ipianlilaliv e mor-
phological cliaraclers were studied 4300
in ca.
specimens
dried from the following herbaria: H.
history of the genus. I.ainank referred
ie 7<",li|
( I I
BRNM.
BM, COI-WIITK.
BC, BCF, BR,
C.
K. VI. llie helerocarpic and homoeaipie species as "ai
caz1. ch. c/l hvk. iran. aca.
jk. k. i.au, niques" and Neckc
""doronics.*" respectively.
i. .1
macb,
linn, ma. maf, mo, ny, ro.
it:, a. s, (1790) even proposed the different genus Aronicur,
i
W
SANT, UPS. W. ZA. and Sanehez-Pedraja
the
L
.
had wide acceptance
the 19th
in
t
photographs and photocopies of specimens, or ad- 1838; Koch.
dolle,
II
ditional information, were available: AY. B-W. rhenbaeh. 1854: Srhur. I860: Ardoino. 1867:
ORM.
B0L0, C-BOIS. C-DC. OK. some
BP. C. CI.. monkai. 1886). although authors
I.I).
<
NAP
P-HA. PI, A. .KB. PAI, and SZB. A of recognized Aronicum as a section of Doroni
list
I
h
1831: Willkomn. hinge. 1870; Nv- eio« arpy was clam Iikova. 1961). In
>rosi. <X
'•
1879; Hoffmann, 1892; Beck, 1893). This the Flora of the H.S.S.R., the classification pro-
l.entham looker posed Willkonnn and Lange was merged
persisted by (18.70)
utilil
,iii(l
i I
iloT.Kii pkii-.'il \ronicum as a sMionym of Doroni- with that of Cavillier (191 1) to distinguish two sec-
cum. Tausch also recognized these two tions Doronicum (Corsehkoya. 1061): section
(1828.) in
groups hut referred the heleroi ar|iic specie.-. lo /\/r- \roniciim Iser. [llano, ser. Carpatica). and section
',,,.
dalianches. not Aronicum. hc\ (ser. \ustriacd. ser. ('.,;,:,.',..:;..
Cassini (1817) estahlislied the genus Grammar- Macrophylla. sei. Parddtinncbcs. and ser. I'lan-
ser.
More
with two species. 6. hill^uldlnm and ,<™r- ia^inca). re.enlly. Fdnioiidson (1978) fol-
lliroti (V.
now subsumed within Doronicum lowed Cavillier classification (1911) instead of
pioides. (Jac<|iiiii. s
monotypic
1773; Lamarck. 1786); de Candolle (1836) de- Corschkova's (1961). but described the
&
scrihed the monotypic genu- l-'ullartonui kd- subsection Isaurica \D. cacdllljolium Boiss.
(l\
maont'iisis). now /A kdnuionense (DC.) Alv. Fern. lleldr.) within section Doronicum. which corre-
(Alvarez Fernandez. 2001). In 1838. de Candolle sponded to Cavillier** section Doronicitstrum
proposed two sections within Doronicum: section (1911).
species The phylogenetic analysis herein confirms what
in luding SOIIH i.l Si net in.
and the monotypic section Chromochaeta. with Do- a preliminary morphological study suggested: the
DC,
ronicum now Senecio morphological characters used are loo labile
llnijolium (Wall.) also in pi,,-
|,>
(Maguire, 1943).
Webb in Webb and Berthelot (1846) treated the (\l\arez Fernandez et ah. 2001). The molecular
genus Perivallis D. Don (in Sweet. 1833-1833: lah. data Irom nuclear ribosomal and chloropla-l >\ \
I
some
228) including live species as a section ol Doion suggest groups but without ei gh support to
One
nmend
iriim (D. cruenlum. D. cchiiialum. pap\ru< inn. formal laxonomic groupings. ex-
I), <
D. tussildfiinis. and D. webbii), but is now treated cepti on is a Meditei group of species i/>.
it
original rank (Nordenstam. 1978). plantagineum group), which receives good support
at its
Cavillier (1907. 1911) divided his study of the from both molecular and morphological data. To
complex laxonomic
genus Doronicum. the hist devoted to the study of a\oi( adding to the al ready hls-
1
llie In. inoi arpic species (1007). and the second to lor\ of the genus, a formal infrageneric treatment
the heleiiM aipic ones I0||). However, he later proposed here but deferred against
,-
.1
. >
i i
concluded (1911) thai llus character was not useful
lo delimit natural groups. Cavillier proposed a new As already mentioned in the introduction, after
classification (1911) that included 3 sections. 7 Cavillieis revision and until the present work, a
subsections, and 3 species w Doronicum: sec- large number of laxonomic action- were taken with-
M
I i 1 1
These new
tion Doronicnslrum i-iil.-ccl. Corsica, suhseet. \us- in Doronicum. include llie dc-ci iplion ol
Fdmondson,
suhseet. Cdrdlophvlld. suhseet. Mdcroplni- species (Sergicvskaja. 1949; 1973.
Iritico.
suhseet. Pardaliaiichia. suhseet. Plantugined. 1978; Chen. 1998. among others) and one subsec-
ld.
(Fdmondson, and
and subsecl. Gnindijlora). section Souliedstrum [D. tion 1978), a few lectotypifica-
&
Jams
Maxim. and sei lion tions (Chacon. 1987; Perez et ah. 1997;
",-,, //,
• • i.
hookeri C. B. Clarke ex Hook.). However, this Tin-land. 1998). Despite all of these actions the ge-
(I),
nus was lacking nomenclatural Thus.
stability.
still
Hookerostrum was described on the basis of a spe- during this study and immediately preceding this
cies limn another tribe {Xannoglotti.s hookeri. \s- work, several nonien. lalural and laxonomic clari-
&
tereae) and the phylogeuetic position (Alvarez Fer- fications were done (Alvarez Fernandez Nieto
nandez 2001) of section Soul rum's only Feliner, 1997, 1999, 2000; Alvarez Fernandez,
el al.. 'least
The 16 names Doron-
taxon [irecludcs rccoguilion the sectional le\el 2001). lectotypification of of
al
&
without splilling the hulk genus. icum in current use (Alvarez Fernandez Nieto
arlilicially ol llie
The subsections Doronicum were defined (Cav- feliner. 1000) gave lhe genu- iiomenelaliiral sta-
in
1911) mainly from the shape of basal leaves. bility leading up to this revision. In llu- work, ad-
illier,
names
the size of leaves, and the presence of cypselae ditional lectotypification of seven belonging
dimorphism. Phylogenetic study of the genus (Al- in Doronicum. although not in current use. is also
varez Fernandez ah. 2001) concluded that these presented lo consolidate and clarify as far as pos-
et
and nomenclature genus. Despite
are not synaponioi pliic characters, llicrclorc sible the of the llie
made
Ca\illicr*s classificalion does not recognize natural cilotls lo locale lypc material for all the
groups.
F\en alter CavillieFs work the relevance of het- cdrpdticum. D. clu-.ii. D. cor>it urn. and D. nriculah-)
of Iberian peninsula (except the Pyrenees)
(lie
occupied by D. carpet a nam, under which four
si
species are recognized.
Other alpine to subalpine species in central
t
Half of the 26 recognized species of Doronicum
Kumpe
arc distributed in and North Africa. Se\en The remaining in Kiiropean species arc emit
tv
ics. D. cataraetaru Austrian Alps and
lrat|. Caucasus, and Turkey and () species are dis- corsieum in Corsic, and both occur in subalj:
). i.
trihulcd in central Asia (Turkistan. Allay. Tibet. habitats.
Yiuuian. and Himalayas). The genus
tin- Doronii urn southwestern Asii
in
i
m
With the exception of Doronicum orientate, each represented by sevi specie». Only one. D. obi
species is restricted to our ol die three well-del mi- gifolium (from the Ca.icas, morphologic;
is), is
Kumpe.
southwestern
lled areas: \sia. central \sia. .juile different from the others. Three of them ar
Doionicnm oiienlalr is distributed in Kumpe and widely distributed: D. macropi, \ilniii (Lain asiis am
southwestern Asia, abundantly the eastern Med- northern Turkey (Caucasus am
in D. doliclioli iclitim
).
iterranean (Orecec. western Turkey, southern south Caspian and ma.ximuin
Italy. of the %-al. (cast
I >.
and Lebanon Syria), and scattered in central Ku- ern Turkey, and south the Caspian Sea). Of lb
ol
mpe. where lis pio\iiiiil\ |o inhabited plaees sug- three remaining species a limited number of spec
gests possibly hay in» escaped from gardens. Delim- imens are known, and this results in a scatterei
iting the natural areas ol distribution also distribution.
is
difficult in the ease ol two other Kllinpeail species; All of the central Asian species (D. altaicum, D
plantagmeum
I), and D. pardalianches. Both were liiii/ueiii. D. jalconeri. gansuen.se. D. kaina
I).
onense. D. slenoglossum) overlap at least in tun
17H7)aml now
(IVna. 1571; Miller. an- considered point of their ion-.
— >ul
il
< I i 1 1
Kingdom
alien plants the United (Harron.
in 19K(>;
&
Clement
Foster, L994).
The
species discussed above
orienlale.
(I), I).
plain, igincmn. and D. paidalianches) occur sim-
in
mountain
regions. but mil in high habitat- lllie up- icrbs. The shape and structure of tin
per Doionicnm hunnai
tree-line). icum. oeeiipying
Kumpe.
- In habitats in Kasleru can he consid- any one. These characters are use-
tt»
ered plantagineum
vicarianl with I), in this region. idispensahle, to discriminate be-
Doionicnm anslriui urn i- wiileb distributed in In tween species. There are fleshy or woody (or some-
rope, most abundantly in the \ustrian Alps. Mac- what woody) rlii/omes in Doronicum. This cliaraclei
edonia, and Ukraine, always in subalpine regions, was described by Cavillier (1911: 199). who pro-
ami
in the Iberian peninsula occurs only in a lew filed histological diagrams, a- ""lubereiix" and
il
To
"lion liihereux." respectively. dislingiii-h be-
K.i- tween these, obsery ations on fresh material are
re-
Doronicum when
by gnindiflai urn. although pressed. Meshy rhizome-
t|lliret|.
The
of these species
iale.
first
When
stributetl of them. is abun- inal more or less terete shape. fresh, fleshy
It
I'yrenees, and the Canla-
in
ones are fibrous and lough, fleshy rhizomes
art
some
ea-ib reeogni/etl Kiiropean species
in (e.g..
D. p/oiilagiiieum. pardalianches, hungari-
I), I),
cum). while woody rhizomes are well represented
Doronicum
he \lps (briquet. I'XHl. cliisu pre- Asian species mucroplnllum. maxi-
is in (e.g..
I), It.
en! in the Alps and !aipalluaus. while D. glaciate mum, D. steiioglossum). In a few cases, rhizomes
I
s restricted to the Alps (mainly the Austrian \lps). are Meshy to somewhat woody and cannot be as-
and brown when
signed hi cliici Ivnc g IK n liilijlonilll. I), ca- fresh, pale \ellow In dry. (.cnotallv
li l;i
,
some
stems are straight, hut zigzag stems occur in
Within a species, rhizome internodes may have species (e.g.. IK a ihi< mil file slems are often
list
).
When
roughly conslanl length and width (e.g.. /A «//«/'- simple and end in a single eapitulum. hear-
ch/h. Fig. K,). or may vary length ami width, ing several capilula. stem branched only
In is in
it)
-
I
resulting in stolon-like structures. Most species the upper part. Fxceptions are seen in stcnog-
I),
have the former condition, and only oricntalc
I),
planlagincum. panlalianchcs. and
(Fig. FA). I). I). branches on the lower part of the stem.
sometimes hungaricum
I),
li
natures Further capilula. any. are on
ter-
first. if
-
-
nodes have brown-tinted
n In-i mini lateral branches, which lor the most part
scale- reiuaiiiing from the sheath of hasal lca\es tvertop the main head. Fach species generally has
from previous years (e.g.. carpctanum. Fig. IF). number one
I), characteristic capilula.
of e.g.. in
I).
i
The
persistent remains of hasal fibers from old men. up panlalianchcs. and more
at<; to ~> in
I),
example, Doronicum uhlnn
petioles occur, for in
han 5 (up to 20) in D. corsicum.
gifolium. hut they are frequcnth absent (e.g..
I).
austnacum.
Fig. ic).
Hyaline, shiny, and smooth trichomes are some-
times present on the younger Ic- of rhizomes and
Leaves simple Leaf.
are
.
Sometimes
also the axils of basal leaves. thc-c
in
used taxonomy
ally in the of the
trichomes are long, abundant, entangled, and while
genus (Cavillier. 191 I), but their usefulness is lim-
to yellow, and they can cover a large part of the
The shape and
ited to the specific level. size of
rhizome onm/alc). Such rhizomes were
re-
le.u.. I), specimen
even
leaves arc variable within single
a
"eriopode" by
ferred as Cavillier (191 199) in
to
1:
some
species. For reason, basal leaves
for
lliis
mnopode"
contrast rhizomes, which lack llu-
to
"'gv
mam
indumentum
(o/umnae). cases
leg.. In
I),
it Sim
leaves necessary descriptive purposes.
for
.ii\-
-cc trichomes on rhizomes, because
difficult to
is
they are short and scarce and can be covered with
as lower, middle, and upper, inserted the
i.e.. in
remains carpctanum, gla-
leaf (e.g..
I), I), clii.sii. I),
basal, middle, and upper third of the stem, respec-
and D. grandiflorum).
ciale,
some species, basal and lower cauline
tively. In
Buds on some rhizomes
are evident fleshy (e.g..
hungaricum. These stem buds can be
D. Fig. Mi).
Basal leaves are petiolate. the petiole being short
seen plants two years or older, but these must
in
much
and wide (e.g.. IK hrnpiclii). or longer than
Sometimes
be collected carefully. ihe scales that
the leaf blade (e.g., columnac). In species with
I),
cover young finds can also be observed.
conspicuous
large basal leaves, sheaths are (e.g..
Two species each have unique rhizomes. Doron-
maximum,
macrophyllum. and
dolicholri-
I), I), I),
" -'- >'• -'.'/.-(.' iiiMinbi >
>,,, .i ii
. • I! i - i I
chum). Aeropetally. along the stem, the petiole
nodes, alternating
lo pi
ii-i
ii .1 li
- ii i i ii
gradually shortens, often leading lo fiddle-shaped
with nodal constrictions., sometimes covered by a
leaves. The upper cauline leaves are reduced, ses-
second seen most
fibrous net. In the type, in
I).
on convex
> in is ins. ited a
-.it
> mi ii
marked
swollen woodv surface. Sometimes, pieces of a i
crophyllum. and panlalianchcs
I),
stem were
dicular the also
to
" ii
i
orbicular, ovate
collected. Although the whole structure has not
The
ovate, as well as fiddle-shaped or bract-like.
leaves can be cordate 2 subeord.ile.
l.a-c (Fig.
ol \).
woodv rhizome, but further study of the subterra-
truncate, or attenuate (Fig. T\. Leaf mar-
nean organ needed. (,, F. I).
is
sometimes
Doronicum gins are generally entire to subentire.
\dventilious roots are present
in
markedly dentate (e.g.. IK cacaliifolium. colum-
and sometimes I),
stenogl.os.sum in D.
and
nac, corsicum. grandiflorum).
I), I),
n
:.!,-
Number
and arrangement of cauline leaves de-
gan. suggesting
t
termine to a large extent the architecture of the
specie- the uuinbei cauline leaves
plant. In -(line ol
and con-
low oricntalc) they are
is (2 to 4. e.g.. I),
fined to the basal third of the stem. In other leafy
icum. or allai-
I),
Afv
—
v Hhizomrs Doronirum. Doronirum onmlolr Imm Ming
in 3513. Doronirm
1. \. (.Ir.-mn H). H.
II
—
Doronirum ratamrlurum (dmwn
from Hopflingrr Doronirum columnar (dmwn
s.n.. I?\1|. K. fnmi
>. N/rj
Number
Volume
3
90,
—
MA).
Doronicum pardalianches (drawn from Rivas-Goda\ A. Basal leaf,
gine 2. A, B. s.n.. —
&
Upper
Doronicum dolichotnchum (drawn from Davis Hedge 29493, K). C.
sal leaf. C. D. —
/W
Dnmmeum
mentum of upper eauline leal. K. '. hnquelii (drawn from 22.S80. E). K. I
I
CM.
IndnmeMliiiii upon eaiiline leaf. Dnnmirnm allairum (drawn froni Krasnoborov etui. <>
ol
middle
leaves arc usually seen the or liasal
csl at
Doronicum species have hemispheric
radiate,
All
parts of stems.
widely campanulate, hoinochromous ea[)itula
to
5A, E) with yellow or green-yellow corollas.
(Fig.
observed specimens and preferably from
dry
in mm
ily
Capitulum diameter ranges from 8 15
to (e.g..
and
For description categorization
leaves.
liasal its
cm
olium and kamaonen.se) and 7 to 8
I),
Work-
terms proposed by the Leal Architecture
tht'
The
D. falconed and D. cataract arum). recep-
(e.g..
ing Group (1999) are used. Most species have an
convex hemispheric, glabrous or pubes-
tacle to
is
venation vein category
acli riiirous lor first (e.g.,
iin
some-
cent. In fruit, the base of the capitulum is
uhjloium. carpclanum. reliculalunh
'« in
I), I),
times widely turbinate.
more
which secondary and veins are or
tertiary
all
Ray flowers are female with strap-like or narrow-
less equally evident. Pinnate venation for the first
ending
obovate generally
elliptic to slightly rays,
ly
vein category occurs central Asian species. In
in
sometimes
in three or two teeth, entire (e.g.. D. al-
marked.
this type, the tertian veins are not well
ta'uum). Disk Mowers are hermaphrodite, aclino-
morphic, and narrowly funnel-shaped.
are equally prominent and thick (e.g.. allaicum.
I),
herbaceous papery
Phyllaries are to slightly at
nansucnu Lamaonensr.
It
I). I). \'. /.
base some species austriacum) and
the in (e.g..
I),
The acrodromous type of venation for the first vein
arranged 2 or A rows, the outer being wider than
in
category restricted to a European group of spe-
is
the inner. In most species the phyllaries are clearly
hungarirum. plantagi-
cies oricntalc,
{!). I), I),
can almost
shorter than the flowers, but they
rav
I..-
and columnar). Intermediate cases be-
nciim.
I),
equal or even longer than them (e.g.. slcno^los
I),
sum. panlalianches). Phyllaries are ovate-trian-
I),
columnar. and
carpali< urn. IK
I), I).
;•
i gular, ovate-elliptic, or ovate-lanceolate to linear.
and between pinnate and inodromoii- vena
ai
s-. I Tin phyllary apex usually acute, except
is ill I).
on D. and glaciate.
in clusii
I),
LA-
gansucn.se where bears a sessile gland (Fig.
il
margins except
Phvllarv are entire, Iiaus-
('). in I).
sknrrhiii where they are fimbriate. group
slijditlv \
plantagineum.
species oricntalc. Iiun-
of (I), I), I).
he distinguished:
i
(I)
bearing
phvlla s with ciliale maruiiis. thin,
hearing
ie- h 1
-r:i|M.'-.< ...» in
in
- . , I. nd trichomes (0.2-1.5
equidistant
leaves inserted the basal
(2 to 4) in
5E-G).
(Fig.
some Furopcan
third of stem. displayed by
is
ill.- Il
species oricntalc, columnar, plan-
(e.g..
I), I), I),
tagineum).
An "ultaicum" generally single eapilu Some species of the genus have dimorphic eyp-
type: a
(2)
him: mostly stem ami variable number of selae (heterocarpy). evident primarily by the ab-
a leafy a
uniform leaves along the stem length, or at sence of a pappus in ray Mowers. (Apselac wrlhoul
( I I )
lower This present some cen- pappuses are also generally larger than those with
least in its half. is in
Asian as well as Kuropean species (e.g.. a pappus in the same specimen (Fig. f>\. 8). Het-
tral I),
"
altaicum. on eroearpv occurs in other genera of Asteraceae such
Jolt ,
I).
•
.
"macroplnllum" type: several capilula: siein as Scnccio. Crcpis. Krigeron, Leonlodon. and Hrl
(.">) \
branched in the upper third: large cauline leaves rrothrra, as well as in other families. ( '.arvophv I-
These
laceae. Apiaeeae. Poaceae (/oharv. 1950).
bract-like leaves on the upper stern. This is re- morphologic differences serve diller.-nl hiirclioiis.
southwestern Asian species. Hcterotheca which the type of di-
stricted to the In lalifolia. In
A
morphism Doronicum. Yenable and
"corsicum" type: several capitula; a variable is similar to
(4)
± ±
number of uniform leaves (.It) evenly insert- Levin (1985) suggested that the pappose cypselae
ed a oiig the in. This is el a iv dispersed away bv wind, while epappose cyp-
-,(«
.
selae near the mother plant. This double dis-
fall
some species the habit does not correspo persal strategy implies the potential to colonize dif-
In
to these patterns (e.g., D. kamaonense and I), si, ferent habitats (\etiable tK Levin. 1985; Tanowitz
Although
which are sometimes bi.mehi from et al., 1987; Imhert et ah. 1996). plants
il
I
and sometimes intermediate patterns on with incompletely developed pappuses on ray Mo-
base),
D. pardalianches. cacalii/olium).
(e.g., I),
Garden
Missouri Botanical
—
i2<m.
(-id.
k.
,
