K. S.

KHATRI1)
Department of Botany, Tribhuvan University, S. N. Campus, Mahendra Nagar, Kanchanpur,
Nepal

"Cardamine pratensis complex"

in the Asian Territory of the USSR

Keywords
C. ~ymanii, C. dentata, C. pratensis Complex, Arctic, Temperate, Siberia, Asian territory,
Soviet Union
Abstract
KHATRI K. S. (1987): "Cardamine pratensis complex" in the Asian territory of the USSR. --
Folia Geobot. Phytotax., Praha, 22: 263--269. -- The taxonomy of the "Cardamine pratensis
complex" in the Asian territory of the USSR is dealt with. After critical analysis of the materials
stored in various herbaria in Leningrad (LE, LEU) and Moscow (MHA, MW) three species,
C. nymanii GAND., C. dentata SCHULT.,and C. pratensis L. s. str. have been recognised pointing
out their intraspecific variations and geographical distribution within the territory concerned.

This is a n e x t r e m e l y complicated complex u n d e r which seven or more taxa have

been recognised from Europe (LSvKVmT 1956, JONES 1964, URBA~'SKA-WoRYT-
KIEWICZ et LANDOL~ 1974). LSVKVIST (1956) for the first time t r e a t e d seven species
under the complex a r r a n g i n g t h e m in thre e groups on the basis of rhizome morpho-
logy a n d geographical distribution. Of these, five species are d i s t r i b u t e d in the Soviet
Union, however C. m2tttb[oli MORErTI a n d C. rivularis ScgtrR e x t e n d their range to
the Carphathians a n d some n o r t h - w e s t e r n lowlands of their E u r o p e a n t e r r i t o r y
while the r e m a i n i n g three are widely d i s t r i b u t e d circumpolar elements. Of these
C. n y m a n i i GA~D. belongs to the Arctic G r o u p while C. dentata SCHULT. a n d C.
pratensis L. s. str. come u n d e r the T e m p e r a t e Group (cf. LSVKVIST 1956).
The taxonomic history of the complex in relation to Siberian material is grounded in the
works of C. A. MEYER (1831) and LEDEBOUR (1841). C. A. MEYER recognised C. dentata and C.
pratensis as separate species, but LEDEROUR retained only C. pratensis placing C. dentata as its
intraspecific unit. He was followed by others, but MAx~MowIcz (1873) merged an entirely different

') Present address: Department of Botany, Leningrad State University, B-164, Universitet-
skaya Naberezhnaya. Leningrad 199164, USSR
264 FOLIA GEOBOTANICA ET PHYT()TAXON0.M[ICA 22. ]9~7

species C. prorepens FISCH. ex DC. w i t h C. pratensis reducing the complex as var. typica MAXIM.
Most of t h e latter authors accepted C. pratensis as separate species, however t h e y also retained
var. typica for one or more forms of the complex.
O. E. SCBtrLZ (1903) acc .pted C. pratensis in the broad sense and referred the Siberian specimens
to the following t a x a along with t h e main type: subsp, angusti/olia (HooK.) O. E. SeHULZ, vat.
dentata (SCRULT.) NEn~R., vat. palustris WIMM. et GRAB., var. /luitans O. E. SCHULZ, L arctica
O. E. Sc~'~-~z and L parvi/olia WII~M. et GRAB. Other authors also accepted C. pratensis as a poly-
typic conspeeies consisting of var. typlca MAXIM., var. parvi/olia WI~M. et GRAB., var. dentata
(SCHULT.) NEILR., var. pcdustris WIMM. et GRAB. and var. ]luitans O. E. ScmyLz, b u t overlooked
the distinctness of the arctic plants viz, subsp, angusti/olia and f. arctics ( B u s c h 1915, KRYLOV
1931 and others).
Later on ILJINSKIJ (1926) accepted C. dentata as a separate species following LI:SD~-~r (1914)
and also established karyological correlations between them, as 2n ~ 24 and 2n ~ 72 (determined
b y SENJA~rI~OvA-KoR~AOINA) for C. pratensis and C. dentata respectively. Although he considered
the Groenland specimens as a separate taxon, however he did not recognise it from the Soviet
Arctic and this seems to have been placed under C. dentata as f. arctics (cf. ILJIsSKIJ 1926: 367).
Other authors also accepted only these two species placing the arctic t a x o n under either of th.~se
( K o M ~ o v 1929, B u s c h 1939). Ultimately, LSVKVIST (1956) recognised C. nymanii as an arctic
species pointing out its circumpolar distribution including the Soviet Arctic; this t r e a t m e n t
has been followed b y others (SI"ASSKAJA 1972, 1978, KOTOV 1979, BERKUTENKO 1983).
Thus following recent taxonomic treatment and critically examining the herbarium
m a t e r i a l s s t o r e d i n t h e I n s t i t u t e of B o t a n y , L e n i n g r a d ( L E ) , L e n i n g r a d S t a t e U n i v e r -
sity (LEU), Moscow State University (MW), and Principal Botanical Garden,
M o s c o w ( M H A ) t h e s e t h r e e s p e c i e s a r e r e c o g n i s e d f r o m t h e A s i a n p a r t s of t h e S o v i e t
Union. In typical forms distinct morphological characters exist for their identifica-
t i o n , b u t s o m e t i m e s i t b e c o m e s r a t h e r d i f f i c u l t d u e t o a c e r t a i n o v e r l a p p i n g of m o r p h o -
logical variations. In herbarium specimens difficulties are often encountered because
of t h e p o o r p r e s e r v a t i o n of t h e leaflets, e s p e c i a l l y t h o s e of t h e r o s e t t e l e a v e s w h i c h
p r o v i d e a v a l u a b l e c l u e t o t h e i r t a x o n o m y . A n a t o m i c a l s t u d y of t h e f r u i t v a l v e s ,
s e p t a a n d s e e d s w a s a l s o p e r f o r m e d b u t t h e y w e r e f o u n d m o r e o r less u n i f o r m .
Similarly, nectar glands do not show taxonomic variations within the complex;
h o w e v e r , p o l l e n a n a l y s i s w a s f o u n d of t a x o n o m i c v a l u e a t s p e c i e s l e v e l . T h u s t h e
T e m p e r a t e G r o u p is c h a r a c t e r i s e d b y m e d i u m l y r e t i c u l a t e d p o l l e n s b u t t h e a v e r a g e
d i a m e t e r i n C. p r a t e n M s v a r i e s f r o m 2 6 . 1 1 - - 2 8 . 8 0 m i c r o n w h i l e i n C. dentata i t
is 2 9 . 6 0 - - 3 3 . 5 0 m i c r o n . I n t h e A r c t i c G r o u p t h e p o l l e n d i a m e t e r v a r i e s f r o m 3 0 . 0 5 t o
3 3 . 2 7 m i c r o n w h i c h c o m e s n e a r e r t o C. dentata b u t d i f f e r s f r o m i t b y f i n e r r e t i c u l a t i o n .

KEY FOR IDENTIFICATION

1. Leaflets of the rosette and cauline leaves thick, smooth, lustrous, veins embedded, t e r m i n a l
slightly larger or almost similar to laterals; rosette leaves w i t h 4 - - 1 0 pairs of 3 - - 1 0 : 2 - - 7 ram,
oblong-lanceolate to orbieulate, more or less petiolate, entire rarely obscure-dentate, deciduous
leaflets; cauline leaves with 3 - - 7 (10) pairs of pinnate or pinnatisect, persistent, 4--15: 1 - 3 ram,
linear-lanceolate, linear, entire leaflets; stem 5--20 (35) cm high, usually simple; petals 8-- 12.5 m m
long, white-lilac with darker veins; fruits poorly developed . . . . . . C. nymanii GAND.
+. Leaflets of the rosette and cauline leaves thin, often pubescent, veins not embedded, t e r m i n a l
distinctly larger t h a n lateral, leaflets in the rosette leaves petiolate, crenate-dentate or wavy;
in cauline leaves sessile or petiolate; stem 10--50 em high, often branched upwards . . . . . 2.
2. Leaflets of the cauline leaves distinctly petiolate, usually deciduous, 3 - - 9 paired, 7--25:
1.5--10 ram, oblong-ovate, linear-lanceolate, crenate-dentate or entire; in rosettes 3--7 (10)
paired, petiolate, 4--15: 3--15 ram, orbiculate-cordate-ovate, wavy oc ahnost entire; stem 15 to
50 cm high; petals 8-- 17 m m long, white to light lilac with darker veins . . . C. dentata SCHULT.
KHATRI: CARDAMINE PRATENSIS 265

+. Leaflets of the upper cauline leaves sessile, persistent, 1-- 9 paired, 8 --20:1 -- 8 mm, obovate,
ovate, lanceolate to linear, entire or sparsely dentate, lowest cauline leaves more or less similar
to the rosette ones but with a higher number of leaflets; rosette l~,a.ves with 1 - 7 pairs of 4--15:
6--22mm, petiolate, orbiculate-cordate-reniform, crenate-dentate, usually hairy and rough
surfaced leaflets; stem 10--45 cm high; petals 5-- 14 mm long, white, pink to dark lilac . . . . .
C. pratensis L. s. str.

Cardamine pratensis 1. Sp. P1. : 656, 1753

Syn.: C. pratensis L. var. parvi]olia WIMM. et GRAB., C. pratensis L. var. typica MAxI~.,
C. pratensis L. var. pratensis GLEAS.
E v e n in t h e s t r i c t sense i t is a h i g h l y p o l y m o r p h i c species. T a x o n o m i c a l l y , S i b e r i a n
specimens h a v e been t r e a t e d v a r i o u s l y till recently, h o w e v e r t h e p l a n t s passed
b y most a u t h o r s a s var. typica M A x I ~ . a n d var. parvifolia W I ~ M . e t GRnB. belong
t o C. pratensis L. S. str. I n Siberia along with t h e t y p i c a l form, specimens with
smaller leaves a n d flowers are of c o m m o n occurrence. These differ from t h e t y p i c a l
ones b y s m a l l e r leaves, leaflets in u p p e r cauline leaves linear, flowers smaller, 5 - - 7 m m
long, d a r k lilac, as a whole t h e p l a n t s are also smaller. T h u s in some morphological
c h a r a c t e r s a n d d a r k e r cotoured flowers these specimens come n e a r e r to C. rivutaris
SCHUR b u t differ b y h a v i n g t h e lowest cauline leaves m o r e or less similar to r o s e t t e
ones, a n d yellow a n t h e r s . These specimens have been referred t o var. parvifolia
while t h e t y p i c a l ones to var. typica b y m a n y of t h e b o t a n i s t s who have d e a l t with
t h e c o m p l e x from Siberia.
These v a r i a t i o n s m a y be a d a p t a t i o n a l r a t h e r t h a n genetic or geographical, as
such f l u c t u a t i o n s a r e also often o b s e r v e d in o t h e r m e m b e r s of t h e c o m p l e x d e p e n d i n g
on t h e h a b i t a t . The d i m i n u t i o n in v e g e t a t i v e a n d floral p a r t s seems to be correlated
with t h e e d a p h i c f a c t o r s as these are f o u n d in u p l a n d s on open a n d d r y localities
while t h e t y p i c a l p l a n t s p r e d o m i n a t e in l o w l a n d mesic h a b i t a t s . The coloration of
t h e flowers is also c o r r e l a t e d w i t h soil m o i s t u r e conditions as t h e y become d a r k e r
in more d r y s i t u a t i o n s . I n t h e l i t e r a t u r e these t a x a h a v e been used free of t a x o n o m i c
vicissitudes passing o t h e r c o m p o n e n t s which are now a c c e p t e d u n d e r different
species of t h e c o m p l e x . H e r e C. pratensis is t r e a t e d as a s e p a r a t e species b u t in the
absence of well e s t a b l i s h e d correlations, these t a x a do n o t seem w o r t h y of recognition.
Cytologically, i t is well e x p l o r e d in E u r o p e where from diploid to o c t o p l o i d n u m b e r s
(2n = 1 6 - - 5 6 , 60, 64) h a v e been r e p o r t e d . Till now, o n l y a h e x a p l o i d n u m b e r
(2n = 48) is k n o w n from Siberia, however t h e pollen a n a l y s i s shows t h e presence
of umre c h r o m o s o n m l races.
I t is w i d e l y d i s t r i b u t e d in W e s t e r n a n d Central Siberia e x t e n d i n g s o u t h w a r d s
to t h e A l t a i a n d S a j a n m o u n t a i n ranges, to N o r t h e r n Mongolia; in t h e far e a s t e r n
t e r r i t o r i e s it b e c o m e s r a t h e r sparse a n d is confined to some localities of A m u r ,
O k h o t s k a n d U s s u r (Sikhotd-Alin a n d V l a d i v o s t o k 1) e x t e n d i n g to t h e K o r e a n
P e n i n s u l a , S a k h a l i n , K a m c h a t k a , t h e K u r i l I s l a n d s (upto K u n a s h i r Isl.1), t o w a r d s
t h e n o r t h i t e x t e n d s to Central Y a k u t s k , t h e n o r t h e r n t r a n s - B a i k a l Mts., L o w e r
T u n g u s k a , t h e P u t o r a n a P l a t e a u a n d t h e N o r t h e r n Urals.

1) In Kunashir Isl. and around Vladivostok it seems to have" been introduced.

266 F O L I A G E O | ~ O T A N I ( ! A ET P H Y T O T A X O N O M I C A 22, 19~7

(Jardamine dentata SCItULT. Obs. Bot.: 126, 1809

Syn.: C. pratensis L. var. palustri~" WIMH. et GaAB., C. pratensis L. var. dentata (ScHIILT.)
NEILtt., C. palustris (WIMM. et GRAB.) PETERM., C. pratensis L. var /luitans O. E. SCHULZ,
C. pratensis L. var. kurilensis KUDO.
I t is more polymorphic than C. pratensis. Although transitional forms are common
due to free gene exchange, two distinct forms are of usual occurrence, placed by
many authors under C. prt~tensi8 as var. dentata and var. palustris. But at present
C. dentata is well accepted as a separate species hence these may be arranged as
C. dentata var. denh~ta and var. palustris. From Europe these have been placed under
C. palustris (WIMM. et G R A B . ) PETERM. as o~. isophylla PETERM. and ~. heterophylla
PETERM. (el. PETERMANN 1846: 47, L()VKVIST 1956).
The first (var. de~tata) is characterised by ovate, erenate-dentate leaflets of the
cauline leaves which are somewhat similar to those of the rosette ones while in the
second (var. palustri.~) leaflets of the cauline leaves are oblong to lanceolate and
entire while in the rosettes they are orbiculate-ovate and obscure-dentate to ahnost
entire. Both are well represented in Western and Central Siberia but in the far eastern
territories the second predominates. O. E. SCHULZ (1903: 535) recognised var. fluitans
for specimens lacking rosette leaves and having fewer or almost none of the lateral
leaflets in the cauline leaves. Similar plants with normal cauline leaves were classified
as var. kurilensis (ef. KVDO 1922: 113) from the Kuril Islands. These modifications
are caused by certain growth conditions such as damp shaded, inundated or sub-
merged habitats under which any component of the complex ,nay be modified,
hence these taxa do not deserve recognition.
Cytologically, well explored in Europe where from octoploid to dodecaploid or
even highploid numbers (2n = 56--96, 118) have been reported, and from Asian
territory (Kamchatka Pen.) a single number (2n = 70) has been reported, but the
pollen analysis shows the presence of more chromosomal races.
It is widespread in the lowlands of Western and Central Siberia, along C. prate~sis,
but becomes sparse southwards in the Altai and Sajan Mrs. In the fare astern territory
it is rather scarce and is confined to some localities of Amur, Okhotsk and Northern
Sachalin, but becomes usual in the Kuril Islands (extending southwards to Iturup
Isl.), (3ommander and Karagin Islands, Kamchatka up to Karyakia Province.
Northward it extends to Southern Magadan Province, Central Yakutsk along the
basins of the Lena and Aldan, northern trans-Baikal territory to Plateau Putorana
in Southern Taimyr, the subpolar Urals etc. Its range frequently overlaps that of
the Arctic Group towards its northern limits where it becomes diffieult to delimit
the territorial boundary between their ranges; however, it starts vanishing from more
northerly localities while the Arctic Group becomes dominant. The plants from
Anadyr and Magadan territories are exclusively placed under C. nymanii by some
authors but the present examination of the herbarium materials shows that C. dentata
also extends in some localities there.
KHATR[: CARl)AMINE PRATENSlS 267

I?ardamine nymanii GAND. Bull. S(~e. Bot. Fr. 72: 1043, 1925

Nyn.: C. pr(~ten,~'i,~ L. wtr. angustiJolict Hoo~:., ('. prate~sis L. vat. propag~li]era NOR.~IAN,
C. pratensis L. subsp, polemonioides l~owY, ('. pratettsi,~ L. subsp, ang~tstiJolia (HooK.) O. E.
SCHULZ,C. pratensis L. f. arctica O. E. SCHmaLZ,C. pratensi,~' L. f. tenuiJolia LINDMAN, C. dentata
SCHVLI'. f. lapponica LIND~A?,-~

L(JVKVIST (1956) reeognised this species for the Arctic specimens suggesting its
circmnpolar distribution but bhought it was absent from most of the Siberian Arctic
(from the Gulf of Yenissei to Cape Schmidt in Arctic Chukotka). But recent reports
show that it is usual throughout Siberian Arctics as in others. L6VKVIST'S observa-
tions were based on limited material from Siberia, particularly from the collections
of Prof. F. g. KJELLMAN who explored certain areas in Arctic Siberia in the early
eighties of the 19th century; consequently his report coincides with these localities.
But at present the Siberian flora is well explored and there is sufficient authentic
material on C. n y m , e n i i in many herbaria of USSR. The present author's analysis
of the material confirms that it is consistently distributed in the Siberian Arctic.
This species also exhibits polymorphism like other components of the complex.
Usually, two forms are eommon: one with orbieulate leaflets in rosette leaves and
the other with oblong to lanceolate ones. According to L6VKVlST (1956) the former
is of European origin while the second is of American origin. The )~uropean form
predominates in the Scandinavian Arctic but the American form is found only in
a small area over there although it is the only form found in arctic North Anleriea.
However, he did not comment on the distribution in the Soviet Arctic.
The present analysis shows that both forms are remarkable in the Siberian Arctic,
but sometimes these variations are barely distinguishable and it becomes difficult
to ascertain whether specimens are of one or the other form. This difficulty is further
enhanced by the poor preservation of the leaflets of basal leaves in herbarimn speci-
mens. Thus, in the absence of field observations it is difficult to delimit the range of
distribution; however in eastern specimens extending eastwards from the mouth
of the Lena River the oblong-laneeolate form seems to predominate, but occasion-
ally orbieulate leaflets are found even in specimens from Arctic Chukotka. Similarly,
the reports on Alaskan material also show the presence of orbiculate leaflets on
on those plants (ef. HULTs 1968:514).
L6VKVlST (I956: 9:),) regards these variations as slight, due to frequencies of a few
genes but also admits that the shape of the leaflets is easily modified by the soil-
water conditions of the habitat. Similar observations were made by SrASSKAJA
(1972). In Southern Taimyr Peninsula orbicnlate plants are found in sub-alpine
belts of B e t u l a - A l n u s - S a l i x components but the leaflets are thinner than the usual
ones while in northern specimens including those of Severnaya Zenllya and other
islands, leaflets are orbiculate and thicker but slightly dentate and the plants are
also feeble. Such specimens were referred to f. arctica (el. O. E. SCHULZ 1903: 536).
Thus it confirms that not only genetic characters are responsible but the ecological
conditions are equally important for such modifications.
Gytologieally, octoploid to dodeeaploid numbers (2n = 56, 60--90) have been
reported from European arctic regions with a preponderance of oetoploid (2r~ = 04)
number (of. L6VKVlST 1956) but from NE Asia only highploid (2n = 88, 90, e. 100)
268 FOLIA GEOBOTANICA ET PHYTOTAXONOMICA 22, 1 9 8 7

numbers have been reported and there is a similar report on the Alaskan material
(Cf. PACKER ct McPfiERsoN 1974).
It is distributed throughout the Siberian Arctic including Novosibirskie islands,
Wrangel, Severnaya Zemlya, Novaya Zemlya and other islands, southwards it
extends to the Commander Islands, Northern Kamchatka, Southern Magadan,
Central Yakutsk, along the Lena and Aldan basins, Lower Tunguska, Plateau
Putorana, Gulf of Yenissei, Jamal Peninsula, Polar Urals etc.
Although C. pratensis L. subsp, chinensis 0. E. SCHVLZ (cf. 0. E. ScnvLz 1903:
530) has been reported from Shensi Province, China, only these three species are
of wide occurrence in Asia and the report of C. pratensis from Hassora, West Tibet
by HOOKER (1872: 138) seems erroneous, as the Himalayan specimens referred to
C. pratensis belong to an other species C. loxostimonoides O. E. SCHULZ. The type
specimen of subsp, chinensis was also observed by the present author in the Botanical
Museum, West Berlin (B); it seems a highly reduced miniature form of C. pratensis
s. str. but in the absence of sufficient authentic material it cannot be concluded
decisively and it is hoped that additional material will provide a clearer picture:
From North America, also, only these three species are known, but C. pratensis
is considered as introduced, by some authors (cf. FER~ALD 1920, ROLLINS 1981).
Thus regarding the complex as a whole, the present dynamic stage, luxuriance
of taxa and the presence of diploid races suggest that the place of origin was in
Europe and the assumption of L6v~vlsT (1956) that the temperate group in Europe
might have radiated from Asiatic refugia remains unclear in the absence of diploid
races, and the poor representation of the taxa in Asia.
The present distribution of these species suggests that diversification is of great
age and the cytotaxonomical studies of many authors show that C. 19ratensis is
the oldest one among these, a tertiary type as suggested by ILJISrSlaU (1926). The
wide variations in chromosome numbers suggest that polyploidization followed
by hybridization is of great importance for speeiation within the complex. :Further,
direct correlation has been established by many authors between chromosome num-
bers and the moisture content of the habitat, showing that the degree of ploidy
within the complex increases with an increase in soil moisture. Thus, possibly,
C. dentata was developed from C. pratensis through polyploidization followed by
subsequent hybridization during the phases of glaciation when cool moist climates
predominated. C. nymanii might have developed through further hybridizations
during northward migration immediately after the glaciers were regressing. The
preponderance of seed-sterility in C. nymanii also suggests a hybrid origin. This
process of speciation took place over the whole newly conquered area, as suggested
by ILJINSKIJ (1926).

SUMMARY

The taxonomic history of "Cardamine pratensis Complex" in relation to the Asian

territory of the USSg is discussed here. :From the material three species C. nymanii,
C. dentata and G. pratensis s. str. are recognised and a dichotomous key is provided
for their identification. Possible synonymy for each species in relation to the territory
concerned has been covered to clarify taxonomic understanding. Intraspecific
variations and their relationships are pointed out. Similarly, their areas of distribu-
KItATRI: CARDAMINE PRATENSIS 269

tion within the territory are given, based on the literature as well as on examination
of herbarium material stored in Leningrad (LE, LEU) and Moscow (MHA, MW).

LITERATURE CITED
B~:R~VTENXO A. N. (1983): K r e s t o t s v e t n i e K o l y m s k o g o Nagorja. -- Vladivostok, 168 pp.
BI~scH N. A. (1915): Faro. Cruci]erae J u s s . -- I n : F l o r a Sibiriae et Orienti E x t r e m i , 2: 177--272. --
Petrograd.
B u s c h N. A. (1939): Faro. Cruci/erae J u s s . ~ I n : KOMAROV V. L. [red.]: Flora U R S S , 8 : 1 4 to
606. -- Moskva, L e n i n g r a d .
C~OCHmTAKOV A. P. (1985): F l o r a Magadanskoj oblasti. -- Moskva, 396 pp.
F~RNALD M. L. (1920): Some v a r i a t i o n s of Cardamine pratensis in America. -- R h o d o r a , 22:
11--14.
H o o x ~ R J. D. (1872): The Flora of British I n d i a , I -- London, 208 pp.
]-IULT]~N Fo (1968): F l o r a of Alaska and n e i g h b o u r i n g territories. -- Stanford, California, 1008 pp.
H U L ~ N E. (1971): The c i r c u m p o l a r plants, 2. -- Stockholm, 463 pp.
TLJINSKIJ A. P. (1926): On the vegetative r e p r o d u c t i o n and p h y l o g e n y of some species of Carda.
mi*~. -- Bull. J a r d . Prine. U R S S , 25 (4): 363--372.
IM R. J. e t al. (I974): F l o r a Coreana, 2. -- P h y o n g y a n g , 393 pp.
JONES B. M. G. (1964): Cardamine L. -- I n : TuTt~r T. G. et al. [ods.], F l o r a E u r o p a e a , I: 285 to
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l~eceived on 16 J a n u a r y 1986