FURTHER TRANSFERS OF GLANDULAR-PUBESCENT SPECIES FROM CHENOPODIUM SUBG. AMBROSIA TO DYSPHANIA (CHENOPODIACEAE)

2008·10.6 MB·English

by  Sergei L Mosyakin

:
FURTHER
TRANSFERS OF GLANDULAR-PUBESCENT FROM
SPECIES
CHENOPODIUM
TO
AMBROSIA DYSPHANIA (CHENOPODIACEAE)
SUBG.
L
Mosyakin
Sergei Steven Clemants
E.
Kholodny
M.G. of Botany
Institute
Academy
of
Sciences of Ukraine
2 Tereshcher 01601 UKRAINE
)kivska Street Kiev, ooklyn,.
ABSTRACT
new
rhe following corribinations in Dysphania are proposed for 21 species pr
&
D
Simon) Mosyakin
Clemants,
*a (L.E.
&
nandonii Watson Mosyakin
Clemants, D.
stellata andley) Mos;
(S. (St
)
INTRODUCTION
The
genus Chenopodium L. (Chenopodiaceae) in traditional wide circumscription has long been
its sus-
pected to be an unnatural (polyphyletic or paraphyletic) union of taxa belonging to different phylogenetic
many
Consequently, and
lineages. segregate genera infrageneric taxa of various ranks were described and
by
accepted within this group various authors an overview in The genus Dysphania
(see Scott, 1978).
R.
Br.
during turbulent taxonomic and nomenclatural experienced numerous from
history
its also shifts: a single
&
genus of a separate family Dysphaniaceae (Pax Hoffmann
1934) or a representative of Illecebraceae or
Caryophyllaceae— mere
Chenopodium
to a section of (Aellen 1930a, 1930b; see also discussion Eckardt
in:
s.l.
& &
Mabry Behnke
Wilson Mosyakin
1967; 1976; Scott 1978; 1983, 1984; Clemants 2002, and
references
&
The
therein). present article continues a series of our publications (Mosyakin 1993; Mosyakin Clemants
&
1996, 2002; Clemants Mosyakin 2003) on Chenopodium and
segregates.
L.
its
s.l.
&
we
(Mosyakin Clemants
Earlier 2002) justified the transfer of species previously placed in Chenopodium
commonly
Ambrosia and
L. subg. AJ. Scott, called henceforth referred to as glandular chenopods, the
to
new
genus and
Dysphania, validated taxonomic combinations in preparation for the treatment of the genus in
&
Flora oj North America north Mexico (Clemants Mosyakin 2003). At that time we made 18 new combina-
of
including 4 and
tions, 11 at the species level, at the section level, 3 at the level of subsection. Nomenclatural
changes were
essentially limited to taxa occurring in North America (both native and introduced), leaving
number
chenopods
glandular be
a of yet formally transferred Dysphania.
to to
Journal of the Botanical Research Institute of Texas 2(1)
426
an
chenopods within
glandular
The taxonomic and nomenclatural decisions treating
Mosyakin The genus Dysphania circumscribed
were further discussed in the Dr. Sci. dissertation of (2003).
Chenopodiaceae
Chenopodium has been accepted the treatments of
for
include glandular taxa of
so as to all
&
Mos-
(Zhu
Mosyakin and China Gelin,
North America north Mexico (Clemants 2003) Flora of
in Flora oj
of
&
from Chenopodium Dysphania
(D.
Clemants At one additional transfer of a species to
yakin 2003). least
&
&
Lambinon
Lambinon) has been made (Verloove 2006) since then.
Verloove
(Murr)
pdseudomultiflora
genus
Here we new combinations in Dysphania for other species not transferred to that earlier.
validate
However,
We on within Dysphania.
making phylogenetic relationships
from conclusion
here specific
refrain
mor-
herbarium and experience, studies of
our
judging from thorough analysis of relevant literature, field
a
new
we Dysphania the circum-
in
chenopods, firmly believe that
and biogeography glandular
phology of
2002 was confirmed
from Chenopodium Since partly
scription a natural phylogenetic unit distinct s.s. it
is
by molecular phylogenetic studies.
DISCUSSION
between three
confirmed some extent deep phylogenetic split at least
Results of recent molecular studies to a
Chenopodium (Kadereit 2003,
branches containing taxa previously traditionally placed in s.l. et al.
(clades)
named
Chenopodieae
informally
Chenopodium appeared scattered in three lineages I, II,
2005). Species of
s.l.
subgenera
mainly corresponded recognized
and The segregation of species between these three clades to
III.
and Ambrosia respectively; see
Hiitonen, Scott,
Chenopodium (subgenera Chenopodium, Blitum (L.) A.J.
of
I.
s.l.
976) indicated: "In Chenopodieae a close relationship
1978). In particular, Kadereit et (2003, III,
Scott, al. p.
Chenopodium
and
Chenopodium and of the Australian species
of the Eurasian species hotrys Teloxys aristata,
by presence multicelluar glandular hairs (type 8
and supported the of
Dysphania
cristatum glomulifera, is
Simon These
By they into subgenus Ambrosia (Scott 1978; 1996). <. .>
in Carolin 1983). that character,
fit
.
Ambrina
Chenopodium
Mosyakin and Clemants subg.
(2002) transfer [sic!
confirm the proposal of to
results
&
- and Chenopodiaceae Kadereit
M. C] Dysphania .>." In a study of the origin age of Australian
to <.
S. S.
.
Mosyakin and Clemants used morphological characters
(2002)
indicated that "[recently,
(2005, 74)
et al. p.
view sup-
Ambrosia from Chenopodium, and included in Dysphania. This last is strongly
separate subg. it
to
by
ported our
results."
would be recognize several gen-
our wide circumscription of Dysphania to
Other
options alternative to
= Moq., or also other
Chenopodium
example, Moq. (including only aristatum L. T. aristata (L.)
Teloxys
era, for
may
American which be recognized either in the original
Moq. an group,
(mainly
Roubieva
related taxa),
American Australian group previously
Moquin-Tandon with addition of other taxa), the
or
sense of (1834),
circumscription.
and Australian Dysphania in traditional
Chenopodium Orthosporum its
placed in R.Br.,
sect.
seem
schemes Chenopodium
of
These groups, corresponding mainly to sections in various classification s.l,
knowledge, has
our present of
such groups genera, level
be However, recognition of as at least at
natural.
to
many
our
In opinion,
paraphyly uncertain phylogenetic positions of taxa.
or yet
negative aspects related to
its
but excluding other
Orthosporum,
expanded circumscription olDysphania (including related sect.
a slightly
morphological
applicable
would be inconvenient, because will erase practically
non-Australian taxa) also
it
broader genus from other glandular taxa.
distinguishing the resulting
diagnostic characters
we
place
For avoiding such problems and for maintaining the nomenclatural stability strongly prefer to
and by
which by glandular trichomes also other
chenopods Dysphania, characterized
glandular in its
is
all
Carolin 1983;
morphological and biochemical characters extensively discussed in literature (Scott 1978;
Simon Bonzani 2003
Wilson Palomino 1990; Schwarzova 1993; 1995, 1996; et al. etc.).
1983; 1984;
et
al.
from
we new nomenclatural combinations the species level resulting
21
at
In the present validate
article
A
narrow concept applied,
moderately species
Dysphania. is
glandular-pubescent species
the transfers of to
and
chenopods"
species of
which mainly with prevalent taxonomic treatments of "glandular
consistent
is
and
We many considered groups are in need of further revisions
Chenopodium admit species of the
that
s.l.
new Dysphania
making combinations in for
more but should not preclude available
that
detailed studies,
New
Mosyakin and Clemants, combinations Dysphania
in 427
them. In addition to our experience and herbarium studies of these taxa, we especially consulted with and
upon
taxonomic by
relied the basic publications Aellen (1930a, 1930b, 1933, Aellen and
1973), Just (1943),
Wilson Simon
(1983, 1984, 1987), (1995, 1996, 1997), as well as other relevant works (Voroshilov 1942;
Weber
Scott 1978: Carolin 1983; 1985; Palomino 1990; Kiihn 1993; Schwarzova 1993; Uotila
et 1997,
al.
2001).
NEW
HE COMBINATIONS
&
Dysphania
Orthospora
Mosyakin
1. sect. (R. Br.) Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002.
w
names
Several of putative hybrids between Australian taxa belonging Dysphania
Orthospora
to sect.
published Wilson
(Aellen 1933, 1961; 1983, 1984). However, since taxonomic remains
their status uncert,
and their hybrid origin mostly in need of confirmation, we refrain from making here any new combii
is
tions for hybrids.
&
Dysphania melanocarpa
Q.M. Mosyakin comb.
Black) Clemants,
nov.
Chenopodium
Basionym: carinatum
R.
Br.
&
Dysphania
pusilla (Hook, Mosyakin Clemants, comb.
nov.
f.) Bash
&
Dysphania
truncata Mosyakin
(EG. Wilson) Clemants, comb,
nc
&
Dysphania
Mosyakin
saxatilis (EG. Wilson) Clemants, comb,
i
The
species was described by Wilson from Western Aus
last (1983)
noted would
species that "[superficially species appear be
this
to
i
genus Dysphania
D. rhadinostachyd) but in seed character
(e.g.
Chenopodium and
to pumilio of the section Orthospoi
allies
its
since the other species of this section have flowers arranged in compact glomerules, have one stamen, and
normally a 5-merous perianth." (Wilson Wilson compared with members
1983, 180). also his species
p.
of the Chenopodium botrys group (which evidently differ from in having 5-merous perianth and
C. saxatilis
and
horizontal seeds) section Margaritaria (which has whitish subglobose, as well as glandular, hairs, and
some
other Brenan and
concluded
distinctive characters, see 1956), that appears that C. saxatile does not
"it
fit satisfactory into any of the presently recognized groups." E.G. Wilson refrained from describing a new
monotypic and expanded
section instead the circumscription Chenopodium Orthosporum
of include
sect. to
we
Fending
his species. further research, keep the species in Dysphania Orthospora.
sect.
&
Dysphania
Adenois
Mosyakin
2. (Moq.) Clemants, Ukrayins'k. Zhurn. 59:382. 2002.
sect. Bot.
&
&
In our previous publications (Mosyakin Clemants 2002; Clemants Mosyakin 2003; Zhu Gelin
et
al.
we
2003) mentioned that opinions vary widely regarding the proper species delimitation in with
this section,
some
authors recognizing just a few species, including extremely polymorphic Chenopodium
ambrosioides
L.
with more
infraspecific taxa, while other authors preferring several (up to 12) narrowly circumscribed species
(Aellen 1929, 1961, 1973; Aellen &Just 1943; Voroshilov 1942; Simon 1996, 1997; Giusti 1997). Sometimes
same
the author (notably Aellen) in different periods of his work was of differing opinions regarding the
We
number
of species related to C. ambrosioides. believe that taxa in this group differ from each other not
many
than
less generally recognized species of Chenopodium Chenopodium),
(subg. especially in the
C.
s.l.
Numerous
album
group. were and some
recognized
infraspecific entities in C. ambrosioides other variable
we
taxa. In the present article are concerned with the species-level taxonomy, and thus for the time being
new
from making any and
refrain infraspecific infrageneric combinations.
Research of Texas
Journal of the Botanical Institute 2(1)
&
comb.
Dysphania dunosa Simon) Mosyakin Clemants, nov.
(L.E.
&
comb. no\
Dysphania Mosyakin Clemants,
oblanceolata
(Speg.)
&
comb.
Dysphania retusa ex Moq.) Mosyakin Clemants, nov. Basionym: Chenopodium retusumjusi
(Juss.
&
Ac
comb. sooanum
Dysphania sooana Mosyakin Clemants, nov. Chenopodium Aellen,
Basionym:
(Aellen)
&
Dysphania tomentosa Mosyakin Clemants, comb. nov. chenopodium tomentosum
(Thouars) Basionym:
&
nov
comb,
Dysphania venturii (Aellen) Mosyakin Clemants, Basionym: chenopodium ambrosioides L. sl
&
Zhurn.
Chenopodium Aellen ex Vorosch., Bot. S.S.S.F
Aellen in Aellen Just, Amer. Midi. Naturalist 30:5 1. 1943. venturii
&
Zhurn. 59:382. 2002.
Dysphania Roubieva Mosyakin Clemants, Ukrayins'k. Bot.
(Moq.)
3. sect.
&
comb.
Dysphania bonariensis (Hook, Mosyakin Clemants, nov. Basionym: Roubievabonarien
f.)
1934.
Pflanzenfam. 16c:491.
comb, nonTen. 1833. Chenopodium haumanii Ulbr. in Engl. GrPrantl, Nat.
1923,
illeg.,
&
comb. .AnalesUni
Dysphania microcarpa Mosyakin Clemants, nov. Basionym: Roubieva microcarpa Phil.
(Phil.)
&
2002.
Zhurn. 59:383.
Mosyakin Clemants, Ukrayins'k. Bot.
Dysphania Botryoides Mey.)
(C.A.
4. sect.
&
&
Mosyakin
Botrys
Dysphania Botryoides Mosyakin Clemants subsect. (Aellen lljin)
Mey.)
(C.A.
4a. sect.
&
2002.
Zhurn. 59:383.
Clemants, Ukrayins'k. Bot.
combination Che-
name Chenopodium often incorrectly cited as a
The subsection within the genus
is
of this
&
name by Koch
published illegitimate,
Koch) Aellen Since the is
nopodium
subsect. Botrys (W.D.J. lljin.
who
only
name and provided description in
should be Aellen its
above subsectional attributed to lljin,
the
which
by Koch,
and
description
pre-1935 publication
Russian, but also included a reference to the earlier
makes
published.
subsection validly
their
&
Dysphania nepaknsis Mosyakin Clemants, comb. nov. Chenopodium nepalense Colla, Herb. Pede-
Basionym:
(Colla)
DC,
Moq. 1849.
Chenopodium in Prodr. 13(2):75.
multiflorum
but
in
Asian morphologically resembles D. schraderiana in its habit, differs
This little-known Central species
having weakly keeled perianth segments with simple non-glandular hairs at the keel (Uotila 1993, 1997).
from Chenopodium Dysphania
was to
recently transferred
Another D. schraderiana
species closely related to
&
Lambinon Chenopodium Schrad. subsp. pseudomultiflorum
foetidum
(Murr) Verloove (=
as D. pseudomultiflora
(Murr) Uotila)
pseudomultiflorum
(Murr) Aellen,
pseudomultiflorum C.
Murr, schraderianum Schult. var.
C.
&
from
South African origin differs
Lambinon indicated that this species of
2006). Uotila (2001)
(Verloove
New
Mosyakin and Clemants, combinations Dysphania
in
&
Dysphania
procera ex Moq.) Mosyakin
(Hochst. Cler
Taxa unplaced Dysphania
5. yet in sections of
&
Dysphania
Mosyakin comb.
stellata (Standley) Clemants,
nov.
Basionym: Chenopodium stdlatumS. Watson,
Proc.
This morphologically deviant species having 6-8 perianth segments (described from Coahuila, Mexico
by and
placed Standley genus Meiomeria then Chenopodium
(1916) in a separate Standley, transferred
to
A
Meiomeria
(Standley) Scott (Scott 1978). Relationships of this little-known taxon with other repres.
J.
Dysphania remain known.
tives of insufficiently
ACKNOWLEDGMENTS
This and our previous on Dysphania were during our work on
article publications initiated treatments of
Chenopodium and Dysphania North America (FN and China and
the Flora of A) Flora of (FC)
for projects,
during several visits of the author to the Missouri Botanical Garden, Louis (MO) and the Brooklyn
first St.
NY
Botanic Garden, The and
(BKL). authors are grateful to the staff of these institutions herbaria for their
FNA
FC
kind help and cooperation. Special thanks are due to colleagues involved in the and projects (listed
Hoch
Nancy Morin
alphabetically): Ihsan Al-Shehbaz (MO), Peter (MO), (FNA, Point Arena,
California),
who
Peter Raven (MO), James Solomon (MO), Nicholas Turland (MO), and James Zarucci (MO),
facilitated
We
Thomas
our research. are also grateful to Borsch (Bonn, Germany), Helmut Freitag (Kassel, Germany),
Gudrun
Kadereit (Mainz, Germany), Aleksandr Sukhorukov (Moscow, Zoya Tsymbalyuk
Russia), (Kiev,
whom
we
and with
Ukraine), Uotila (Helsinki, Finland), other colleagues discussed various
Pertti issues
related taxonomy, morphology and geography Chenopodiaceae. Thanks due Barney Lipscomb,
to of are
to
Donald and two anonymous comments on
reviewers manuscript.
Pratt, for their valuable the
REFERENCES
Journal of the Botanical Research Institute of Texas 2(1)
430
63:490-492.
Chenopodium
Jahrb.
930b. neue Sektion der Gattung (Sect. Tetrasepola). Bot. Syst.
Eine
Aellen,
P.
1
Orthosporum der Gattung Chenopodium L Verh. Naturforsch. Ges. Base!.
1933. Die Arten der Sect.
Aellen,
P.
44:308-318.
Lehmann
von Verlag,
961 Chenopodiaceae. G. Hegi. Illustrierte Flora Mitteleuropa. Bd. 3.T. 2. Aufl. 2.
Aellen, In:
P. 1
.
& 533-747.
MQnchen Hamburg.).
Parey Pp.
(Reprinted 1979: Paul Verlag, Berlin
in
und
&
L Ambrina Hook, Nigrescentia
Chenopodium (Spach) Benth. Sect.
von
Formenkreis Sect.
1973.
Aellen,
P.
Hung. -1
Acta Acad. -4):1
Aellen. Bot. Sci. 9(1 2.
1
Chenopodium L Amer.
genus
American the Midi.
Key and synopsis of the species of
andT. 1943.
Aellen, Just.
P.
):47-76.
Naturalist 30(1
MA
the aromatic
and 2003. Morpho-histological studies in
Ariza
Bareoza, Bugatti, Espinar.
Bonzani, G.E. L.
N.E.,
74:207-225.
Chenopodium from
Argentina. Fitoterapia
species of
Kew
[Volume
new genus Chenopodium from of 1956.]
A
section of the Africa. Bull.
1956.
Brenan, P.M.
J.
11:165-167.
103:451-466.
Chenopodiaceae and Amaranthaceae. Jahrb.
Bot. Syst.
1983.The trichomes of the
Carolin, R.C.
America
Chenopodium North
Dysphania Brown; Linnaeus. Flora of
and 2003. In:
Clemants, Si. Mosyakin. R.
S.E.
New
York
North America north of Mexico. Oxford University Press,
Committee, of Vol. 4.
eds. Flora
Editorial
&
267-299.
Oxford.
Pp.
Chenopodium und Bauhinia 3:327-344.
von Dysphania mil mit illecebraceae.
1967.Vergleich
EckardtJ.
Cordoba.
Conicet,
fanerogamica Argentina. Fasciculo40. Proflora,
Chenopodiaceae.
Flora
1997.
Giusn, In:
L.
Amaranthaceae and Chenopodiaceae and the
Phylogeny
and 2003. of
Borsch, Weising, H. Freitag.
Kadereit, G.,T. K.
C 164:959-986.
-photosynthesis.
evolution of Sci.
Intl. J. PI.
4
Chenopodiaceae. Organisms,
and age
and 2005. Origin of Australian
Gotzek, Jacobs, H. Freitag.
Kadereit, G., D. S.
&
5:59-80.
Diversity Evol.
Chenopodiaceae.
and
RG. Wilson). 993.
Hedge, Uotila,
Kuhn, U. (with additions by V. Bittrich, R. Carolin, H. Freitag, I.C. P. 1
New
253-281.
ed.The Families and Genera of Vascular Plants. Vol. 2. Springer-Verlag, York. Pp.
Kubitzki,
K.
In:
Dysphania
the systematic position of
and sieve-element
and P-type plastids:
976. Betalains
Mabry,TJ. H.-D. Behnke.
1
25:109-1
(Centrospermae).Taxon
11.
R. Br.
nouveaux genres de Chenopodees. Ann. Nat. Bot.
de Sci.
Descriptions plusieurs
Moquin-Tandon, 1834.
A.
1:289-294.
and
An system Chenopodium L (species of Europe, North Central Asia).
Mosyakin, Si. 1993. outline of a for
50:71-77.
Zhurn.
Ukrayins'k. Bot. (Ukr. Bot.
J.)
Chenopodiaceae
L Taxonomy, phytogeography and genesis of the family Vent. Dr. Sc. Dis-
2003.
Mosyakin,
S.
Academy
Kholodny of Botany of the National of Sciences of Ukraine, Kyiv (Kiev). [In
M. Institute
sertation. G.
and summaries.]
Russian
with English
Ukrainian,
L
New Chenopodium (Chenopo-
and combination
and 1996. infrageneric taxa in
Mosyakin, S.L Clemants.
S.E.
Novon
6:398-403.
diaceae).
New (Chenopodiaceae):
Dysphania
nomendatural combinations
and 2002. in R. Br.
Clemants.
Mosyakin,
S.L. S.E.
59:380-385.
Zhurn.
occurring North America. Ukrayins'k. Bot. (Ukr. Bot.
taxa J.)
in
Chenopodium
between and
Cytogenetic Teloxys
and distinction
Mercado. 990.
M.
Palomino, G., Segura, R. Bye, P.
1
SouthW. 35:351-353.
(Chenopodiaceae). Naturalist
Harms. Die naturlichen Pflanzenfamilien. 2 Aufl.
and Hoffmann. 934. Dysphaniaceae. A. Engler, H.
A. In:
Pax, K.
F. 1
&
272-274.
Duncker Hunblot.
Pp.
Vol. 16c. Berlin:
L
Chenopodium
mancher Gattung
und Arten der
karyotaxonomie evolution
zur
Schwarzova, 1993. Beitrag
T.
48:421-428.
und
Ambrina).
Biologia (Bratislava)
(Sektion Botryoides
Chenopodium L and genera (Chenopodiaceae). Bot.
AJ. 1978. A review of the classification of related
Scott,
100:205-220.
Jahrb.
Syst.
L Ambrosia
Chenopodium sous-genre AJ.
du genre
paleobiogeographique
Simon, LE. 1995. Essai sur I'histoire
71:127-142.
Biogeographica
Scott.
New
Mosyakin and Clemants, combinations Dysphania
in
431
Chenopodium L
Simon, LE. 1996. Notas sobre subgen. Ambrosia
AJ. Scott (Chenopodiaceae). 1,Taxonomfa.
2.
Fitogeografia: areas Anales Madrid 54:137-148.
disjuntas. Jard. Bot.
SimCn, LE. 997. Variations des characteres chez Chenopodium subgen. Ambrosia
foliares Adenois (Cheno-
sect.
1
en Amerique du
podiaceae) taxonomique
Sud: valeur Adansonia
et evolutive. 9:293-320.
1
New
Chenopodiaceae.
Standley, RC. 1916. Amer.
FI.21(1):1-93.
N. York.
In;
993.Taxonomic and nomenclatural on Chenopodium
Uotila, notes the Ann.
P. 1 in Flora Iranica area. Bot. Fennici
30:189-194.
Chenopodium. Hedge
Uotila, 1997. et 1997. des Hochlandes und
Flora Iranica: Flora Iranischen der
P. In: I.C. al.
Umrahmenden
Gebirge. Persien, Afghanistan, von West-Pakistan, Nord
Teile Azerbaidjan, Turkmenistan,
Iraq,
ed. K.H. Rechinger. No. 172. Chenopodiaceae. Akademische Druck- 24-59.
u.Verlagsanstalt, Graz. Pp.
Uotila, 2001 .Taxonomic and nomenclatural notes on Chenopodium (Chenopodiaceae)
Ann.
P. for Flora Nordica.
Bot. Fennici 38:95-97.
and The new
Verloove Lambinon. 2006. non-native vascular of Belgium: nothospecies and new
F. J. flora a three
combinations. Geogr. 7-220.
Syst. 76:21
PI.
Voroshilcv Overview Chenopodium L
V.N. [Woroschilov; Voroschilov] 1942. of species of Ambrina
(Spach)
sect.
Hook.
Zhurn.
Bot. 27(3/4):33-47.
S.S.S.R. Russian.]
[In
fil.
W.A. The genus
Weber, 1985. Teloxys (Chenopodiaceae). Phytologia 58:477-478.
A
Wilson, P.G. 1983. taxonomic revision of the tribe Chenopodieae (Chenopodiaceae)
Nuytsia
Australia.
in
4:135-262.
Chenopodiaceae.
Wilson, P.G. 984. George, of Government
1 In: A.S. ed. Flora Australia. Vol. 4. Australian Publish-
81-317.
ing Service, Canberra.
Pp.
Wilson, P.G. 987. Generic status in the Chenopodiaceae. Australian Syst. Bot. Soc. Newsl. 53:78-85.
1
Wu
Zhu
and
Gelin (Chu Ge-ling), S.L Mosyakin, S.E. Clemants. 2003. Chenopodiaceae. Zhengyi and PH. Raven,
In:
eds. Flora of China. Vol. 5 (Ulmaceae through Basellaceae). Science Press, Beijing & Missouri Botanical Garden
351-414.
Press, Louis. Pp.
St.