A Taxonomic Synopsis of the Genus Salsola (Chenopodiaceae) in North America

9 Pages·1996·13.3 MB·

by  Sergei L Mosyakin

TAXONOMIC
A
SYNOPSIS OF
Mosyakirv
Sergei
L.
THE GENUS
SALSOLA
(CHENOPODIACEAE)
IN
NORTH
AMERICA^
Abstract
A
key and synopsis of the genus Salsola L. (Chenopodiaceae) in North America north of Mexico are presented,
A
including the taxonomy and distribution of six recognized species. new combination proposed: Salsola kali L.
is
subsp. pontira Mosyakin.
(Pallas)
The
genus Salsola sensu which includes and Halimione means
L. lalo, Atriplex L. Aellen. that the
It
the segregate genera Caroxyion Thunborg, Clima- generic boundaries and
of Salsola related taxa are
=
coptera Botschantsev, IlypocyUx Woloszczak Dar- in need of a systematic revision. also evident
It is
niella Maire, Neocaspia Tzvelev, Nilrosalsola Tzvel- that Salsola in the traditional sense should be re-
and 130-150
ev, Xylosalsola Tzvelev, comprises garded as a group of genera, rather than as a nat-
species, which are especially numerous in the arid ural genus. The taxonomic problems related
to
and coastal zones of Eurasia, including Central forming a generic concept within group have
this
Asia (all sections of the genus), the Middle East, been discussed recently by Tzvelev (1993). He ac-
and the Mediterranean region. Some species, most- cepted several quite distinct genera, including Ca-
ly representatives of Salsola sect. Salsola (includ- roxylon, which is represented in North America by
ing sect. Kali Dumorlier), also occur in other only one alien species, usually nTerred as 5a/-
to
regions of the world as introduced synanthropic 5o/a t^ermicu/a^a; other North American taxa be-
all
weeds, including in North America. long Salsola sensu
to strlcto.
The considerable biological diversity of arbores- As noted above, species of North American
all
cent and shrubby species oi Salsola sect. Caroxyion Salsola should be regarded as naturalized or casual
(Thunberg) Fenzl (or the separate genus Caroxyion) aliens native Eurasia.
to
is confined also to northern (Mediterranean), south- In the course of preparing a taxonomic treatment
em, and
eastern parts of Africa (see Botschantzev, of the genus for the Flora of North America has
it
Some
1969, 1975a, b). "satellite" genera closely become evident that the taxonomy, nomenclature,
related to Salsola {Acllcnia Ulbrich, Physandra and distribution of some species need
clarification.
&
Botschantzev, Horanlnivia Fischer Meyer, Extensive herbarium from MO, GH, NY,
Gir- collections
US
gensohnia Bunge) are widely recognized by bota- and served as the base for this study. Compar-
nists, while some others are not. However, in most ative Eurasian material from LE, KW, MHA, and
cases the latter segregate genera, usually treated as some other Russian and Ukrainian herbaria was
synonyms Sa/Wa
of Salsola sensu lato or recognized as also examined. Types of R. and
Br.
au,s/ra/JA-
MO BM
infrageneric taxa, not distinct from Salsola caroliniana Wall, sent on loan from
S. to
1
<
sensu stricto, than, for example, such readily ac- were also consulted, as were photographs of Lin-
cepted pairs of genera as Kochla Roth and Bassia naean specimens from LINN,
Allioni, Salicornia L. and Arthrocnemum Moquin- Following a brief synopsis of the Salsola spe-
is
Tandon, Corispermum L. and Anthochlamys Fenzl, cies occurring in North America north of Mexico,
&
thank Clifford W. Cr()in[)ton (Agriculture Agrifood Canada, Ottawa, Ontario. Canada), Noel H. Holmgren (New
I
'
York Botanical Garden, NY, Stanwyn Museum
U.S.A.), C. Shetler (National of Natural History, Smithsonian Institution,
UX
Washington D.C., U.S.A.), Ueila M. Shultz (Utah State University, Logan, U.S.A.), Nikolai N. Tzvelev (Komarov
Botanical Institute, St. Petersburg. Russia), and Stanley L. Welsh (Brigham Young University, Provo, UT, U.S.A.) for a
Amy
productive discussion and valuable comments, and Anne Keats Smith and Scheuler McPherson (both Missouri
MO,
Botanical Garden, St. Louis, U.S.A.) for their help in the preparation of the manuscript. The kind help of the staff
MHA,
of BM, BRY, GH, KW, LE, MO, NY, and US herbaria greatly appreciated. Special thanks are due Nancy
is to
R. Morin, Convening Editor, and tlie staff of the Flora of North America Project (Missouri Botanical Garden, Louis,
St.
MO,
U.S.A.).
^ N. G. Kholoilny Institute of Botany. 2 Tereshchenkivska Kiev, 252601 Ukraine.
Str.,
Ann. Missouri Box. Card. 83: 387-395. 1996.
388 Annals the
of
Garden
Missouri Botanical
with some notes on their taxonomy and distribution. cent or hispid. Stems branched (very rarely simple).
com- ascending Leaves mostly
Unfortunately, impossible present a erect, or prostrate. alter-
to
is
it
plete account of the North American distribution nate (occasionally lower ones subopposite), sessile,
margin
for all Salsola species, mostly (hie to taxonomic semiterete, lanceolate, linear, or filiform,
confusion (especially for species of the 5. kali ag- entire. Inflorescences spicate. Flowers normally bi-
gregate) in the literature. The distribution state- sexual, solitaiy (rarely in twos or threes, but in this
ments in this account are based on relial)ly iden- case lateral flowers poorly developed) in axils of
segments
tified herbarium specimens seen in the course of bracts, with 2 bracteoles. Perianth 5, at
this study. For more or less common species only maturity covering the fruit and in most species de-
membranous
states, provinces, and territories are cited in the vcloping a transverse dorsal or almost
following synopsis, while for the rare aliens more coriaceous wing (sometimes only 2 or 3 perianth
Stamens
detailed information presented. segments winged, or all wingless). 5.
is
This does not pretend solve prob- Styles and stigmas 2 Seeds usually horizontal;
article to all (3).
embrjo
lems concerning distribution and ecology of intro- pericarp adherent; spiral.
duced species o[ Salsola in North Ameriia. Its pri-
The name derived from Latin sal, salt; or sal-
is
mary aim provide a more reliable taxonomic
is to
The name Salsola was used by Ces-
sus, salty. first
background
for further studies.
now known
519-1603)
alpino for a plant that is
(1
Moquin-Tandon
as Halogeton salivas (L.) (basio-
Taxonomy North America
of Salsol\
in
nym:
Salsola saliva pul>lished in 1762).
L.,
TYPE;
Salsola Sp. 222. 1753. Salsola
L., PI. 1:
Vernacular names. Saltwort, Russian-thistle
&
soda by
L. (lectotype, selected Britton
(English); Soude, Salsovie (French); Salzkraut (Ger-
Brown,
1913).
man); Soda Solyanka (Russian), So-
Kali, (Italian);
Annual subshrubs Old World lyanka, Kurai (Ukrainian); Solanka (Polish);
herbs, or the
[in
±
also shrubs and small trees], glabrous or pubes- Slanobyl (Czech).
Kkv to Taxa Noinn Amkkica
AiniKiciAL SAisoiA
in
la. Annual hed)s, glabrous or papillose-hisj)id; leaves and bracts with spinose (or at least niucronulate) apex;
perianth segments completely glabrous, or indistinctly papillose (occasionally ciliate at margin).
2a. Leaves (especially lower ones) mostly opposite, with a[)(^x mucronulale, not spinose; bracts distinctly
swollen base, alternate or subopposite; perianth segments normally with crenate or pectinate-ciliale
at
soda
apex: plants always glabrous 3. S,
2b. Leaves alternate (sometimes only 1-3 pairs of lower ones subopposite), with spinose or spinescenl
all
some
(rarely almost mucronulate) a[>ex; bra(;ts not swollen, or In species indistinctly swollen at base,
normally alternate; perianth segments with acute entire apex (sometimes papillose at margins, but never
crenate or pectinate-ciliate); plants papillose-hispid or occasionally glabrous.
mm
±
Leaves herbarium specimens mostly 1-2 broad, gradually
3a. fleshy (in living plants), linear, in
narrowed into rather firm apical sj)ine; bracts reflexed at maturity.
4a. Perianth segments with long-acuminate or long-subulate spinose apex, at maturity forming a
mm
7-12 open
slender columnar beak above the broad wings; fruiting perianth diam.; plants of
sands and inland saline hal)itats 3. S. paulsenii
4b. Perianth segments with short-acuminate or Iriangidar apex, never forming a columnar beak at
mm
maturity; wings absent or shorter; fruiting perianth 6-7(10) diam. or less; plants of maritime
saline habitats (seashores, tidal marshes, etc.) 1. S. kali
5a. Perianth segments with rigid, subspinose apex and distinct midvein; bracteoles not swollen,
free la. S. kali subsp. kali
-
5b. Perianth segments with weak apex and obscure mi<lvein; braeteoles swollen, connate at least
base lb. kali suhsp. pant ica
5.
at
-
3b. Leaves normally not fleshy (occasionally somewhat fleshy in plants growing in saline and alkaline
mm
hat)itats), narrowly linear to filiform, in herbarium specimens mostly less llian broad, in most
1
weak appressed
cases abruptly narrowed into apical spine (mui-ro); bracts reflexed or at maturity.
6a. Bracts ap[)ressed an<l strongly imbricate at maturity, gradually narrowed into subulate
spinose ap<^x; inflorescence narrowly spicate, rather dense, not interrupted at maturity;
mm)
perianth segments wingless or rarely with narrow (usually less than erose wing;
1
plants normally erect, branched al>ove the base, or with a few slender branches near the
base.. 4. 5. collina
-.
±
6b. Bracts reflexed, not imbricate at maturity, in most cases abruptly narrowed into spinose
or submucronulate apex; inflorescence spicate, at maturity interrupted at least in lower
perianth segments normally with membranous wing; plants erect or ascending,
half;
branched from the base.
Volume Number
3 Mosyakin 389
83,
1996 American
North Salsola
7a. Perianth segments wilh long-acuminale spinose apex, at maturity forming a slender
mm
columnar beak above the wings; fruiting perianth 7-12 diam paulsenii
3. S.
7b. Perianth segments with obtuse to weakly acuminate or reflexed apex, at maturity
mm
not forming a columnar beak; fruiting perianth normally less than 8-10 diam.
tragus
2. S.
1
lb. Perennial subshrubs covered with glabrous (smooth) and minutely denticulate hairs (sometimes becoming
±
glabrous at maturity); leaves and bracts obtuse, without spinose apex; perianth segments with pubescent
(not papillose) apex 6. 5. vermiculata sensu lato
1. Salsola kali L, Sp. 222. 1753. TYPE: la. Salsola kali subsp. kali
PI.: L.
Herb. Burser XVI(2): 24 (lectotype, selected
Stems normally papillose-hispid or
(var. kali)^
&
UPS
by
Jonsell Jarvis (1994), not seen).
rarely glabrous polysarca G. W. Meyer).
(var.
F.
See discussion below. The species described
is
Bracteoles free, not swollen. Perianth segments
from Europe: "Habitat Europae
in litoribus
with almost spinose apex and mid-
rigid, distinct
^?
mans.
vein.
Annual 5-50 cm
herbs, papillose-hispid or
tall, Habitats, Seashores, marshes, sandy places
salt
Stems branched from
glabrous. the base, erect, in coastal regions and other saline maritime habitats,
rarely ascending; branches arcuate or somt^times 0-100
very rarely in ruderal inland habitats; m.
Leaves
prostrate. alternate, linear, fleshy, in her-
Herbarium specimens were
Distribution. ex-
mm
barium specimens mostly 1-2 broad, semiter-
amined from the followiniif states, provinces, and
±
eetiee,, nnoorrmmaallllyy nnoott sswwoolilieenn aati boaassee;; aappeexx z: ggrraadouuaaliliyy
FRANCE
, . *t/-^t- / i
(overseas department):
territories:
A r n St.
narrowed ' ^ ,\ nrm • T •
into rather spine. Inflorescence spi- ^. ^\ ^ ^
,,.
,. .^
, CANA.DA.: New . ,
Pierre Miquelon. Brunswick,
et
cate, interrupted at maturity. Bracts reflexed, not
Newfoundland, Nova Edward
Scotia, Prince Island,
imbricate at maturity, alternate, narrowing into su-
Quebec. U.S.A.: Connecticut, Delaware, Maine,
bulate spinose apex. Flowers normally
solitary in
New New
Maryland, Massachusetts, Hampshire,
reduced upper
axils of bracts or leaves. Perianth
New
Jersey, York, North Carolina, Pennsylvania,
segments glabrous, with weak or firm apex, ma-
at
Rhode
Island, South Carolina, Virginia. This sub-
turity wingless or with comparatively narrow wing
species possibly also occurs in the District of Co-
subsp. pontica sometimes prominently winged),
(in
lumbia, U.S.A. native coastal maritime
It is to
becoming connate and united with bracteole bases,
regions of western and northern Europe and nat-
is
3—5
or bracteoles Fruiting perianth
free. ca. (rarely
many
uralized in coastal regions of the world.
mm
up
diam.
to 8)
lb. Salsola kali subsp. pontica Mo-
L. (Pallas)
&
Chromosome 2n 36
number.
(Bassett
syakin, comb. nov. Basionym: Salsola kali var.
Crompton,
1970).
pontica Pallas, PL: 37. 1803. Salsola
lllustr.
pontica (Pallas) A. Degen, Flora Velebitica 2:
Distribution. Native to maritime coastal areas
TYPE: BM,
47. 1937. authentic specimens
in
of Europe, northern Africa, southwestern Asia;
in-
lectotype not selected; see discussion below.
many
troduced and
naturalized in other coastal
regions of the world, including North America. This
Stems in most cases glabrous glabra Forss-
(var.
=
species represented by two possibly
is (or three) kal Salsola pontica glabra Tzvelev, Ukray-
var.
subspecies. Zhum.
sometimes
ins'k. Bot. 50(1): 82. 1993), or
papillose-hispid pontica). Bracteoles swollen.
(var.
Note on The specimen LINN
lectotypification.
connate base. Perianth segments with weak apex
at
was
315.1 selected as the lectotype o{ Salsola
kali
g^j obscure midvein
by Jafri and Rateeb (1978). This specimen repre-
Habitats. Seashores, marshes, sandy
salt pi
sents a typical form of the species. However,
it
regions, rarely in ruderal inland habitats;
^^^
lacks a Species Plantarum number, and probably '^ ^^f
0-100
m.
was added to the Linnaean collection aftiT 1753.
Because of that this lectotypification was supersed- Herb specimens were
Distribution. ex-
ed by Jonsell and Jarvis (1994) in favor of the Bur- amined from
the following states of the U.S.A.: Al-
ser specimen cited above. The second Linnaean abama, San
California (one locality: Nicolas Island,
LINN
specimen
Identified as S. kali, 315.2, also Lat. 33"15'N; Long. 119°30'W, U.S. Naval Radio-
lacks a Species Plantarum number; moreover, logical Defense Laboratorj^ near road above sand
it
seems be 100 Foreman
to identical to S. collina Pallas. spit at elevation, R. E. (No. 42),
ft.
390 Annals the
of
Garden
Missouri Botanical
E, C. Evans III, S, C. Rainey), Driawarc, District there are any obstacles for selecting a particular
of Columbia, Florida, Georgia, Louisiana, Massa- specimen at BM, the lectotypificatioji could be
chusetts, Marj'land, Mississippi, North Carolina, l)ased on the illustration in Pallas (1803).
New New
Oregon one secondaiy, antliropogenous areas of dis-
Jersey, York, (only locality: In their
on ballast groimds of Albine, Porlland, 29 Sep. tribution (particularly, in North America) both sub-
1910, W. N. Suksilorf South Carolina, Texas, spec-ies of kali are often repn^sent(Ml by conver-
s.n.).
.S.
and Virginia; this subspecies also (H:curs in Mexico geni or deviate forms and seem to be less
and seems to be common in coastal regions of South morphologically and geographically separat<»d from
America. native coastal maritime regions of each other than in their native Eurasian areas (e.g.,
It is to
southern Europr^ noilbem Africa, and southwesteiTi North American specimens of 5. kali suhsp. pontica
Asia, northward to Great Britain, eastward to the often approach S. tragus in having quite broad
common
many
Eur-
Caspian Sea, and locally naturalized in wings, a charact<M- that is not as in
is
Due
coastal regions of the world. asian littoral plants). to this peculiarity of in-
This subspecies closely related to 5. kali troduced plants, would be reasonable for the pur-
is it
abandon some
subsp. kali, and in Eurasia replaces the latter on pose of nomenclatural stability to
names
shores of the Mediterranean, Black. Azov, and unceilain of taxa described within the S. Aa//
TX
fc. y
Caspian nomenclature complicated and aggregate from various parts of the world (including
Seas.
Its is
unresolved. For a long tinu' was known in S, caroliniana and australis; see also discussion
still .S.
it
European under misapplied under
botanical [he Ix^low, tragus),
lit(^rature
.S.
names tragus (sensu auct., non L.) or S. kali Wht^n immature, subspecies />on/R*a is almost in-
*S.
subsp. tragus (L.) Celakovsky (in part, excluding distinguishable from S, kali subsp. kali Unfortu-
the type of the basionym). According to Botschantz- nately, many specimens of 5. kali sensu lato
i
ev (1974), the southern maritime taxon of the S, represented in American lierbarla by hnmature
aggregate conspecifie with caroliniana plants. More detailed studies using fruiting material
kali
is
.S.
Walter (type at BM), which seems to be the earliest are needed to clarify the distribution of both taxa
valid name the species level. Tzvelev (1993), in North America. However. S. kali subsp. pontica
at
however, regarded caroliniana as a synonym of certainly a more southern subspecies (in both
S. is
tragus sensu have studied a small frag- North Americ^a and Europe), and seems to be the
stricto.
S.
I
merit of the type specimen of caroliniana sent as only race of S. kali occurring in littoral habitats
.S.
MO.
a loan to This immature [)lant evidently be- fiom South Carolina to coastal Texas. It is also
How- known Mas-
longs to 5. kali, not to S. tragus sensu stricto. from scattered localities as far north as
New
From Hampshire Newfoundland
ever, impossibh^ to assign with certainty to sachusetts. to
is
it it
any subspecies of Moreover, in American only subspecies kali seems to occur, and is quite
S. kali. it
common
name being gradually
literature the 5. caroliniana, or S. kali var. as far soutli as Virginia,
was sometimes applied replaced southward Virginia and North Caroli-
caroliniana (Walter) Nuttall, (in
other taxa of the kali aggregate. Because of na) by subspecies pontica,
to S.
this uncertainty in the proper identity of S, caroli- Both subspecies of S. kali were probably the first
niana, selected as a basionym for the subspecies taxa of Salsola introduced to North America (evi-
I
the name given by Pallas (1803) the coastal Eur- dcntly, on ship ballasts) shortly after the establish-
to
asian race of the kali aggregate. At least, this nuMit of the fust European settlements and the be-
S,
name refers to the native Eurasian taxon, as was ginning of colonization of the continent.
it
noted by Degen (1937) and Tzvclcv (1993). \^'hen Salsola kali is confined mostly to coastal saline
describing his Salsola kali var. pontica, Pallas habitats; however, rarely occurs as introduced in
it
away
(1803) had his disposal specimens from coastal rud(M-al inland habitats, but usually not far
at
mod-
habitats of the Black Sea in the Crim(^a near from the coast.
em
Chcrsonesi Tauricae
Sevastopol
('\ in littore
. .
crescentes pro distincta specie ha])uissem, nisi 2, Salsola tragus L., Cent. PI. 2: 13. 1756. Sal-
Ice--
Maeotin mediterrancis Tauriae/^ Pal-
tae circa et in
1803: and probably some additional spec- Prodr. Fl. lioehmen 2: 155. 1871 (see discus-
las, 37),
TYPE:
imens from coasts of Black and Azov sion under S. kali subsp. pontica).
a(ljac(*nt tlie
LINN
Seas, when* only on<^ littoral taxon of the 5. kali 315.3 (lectotype, selected here; see also
known and Degen, 1937; Tzvelev, 1993).
aggregate occur. Tlu* description
to
is
illustration in protologue are also diagnostic.
th<^ I
^ ^y^
p^^^,^^ ^^ ^^^^, h^^„ ^.
^^^,^^^,^^ ^^^^^,^^^^/.^ p^^^^j^
among
chance
did not liav(* a to select the lectotype ]«]() (emend. Rotschantzev. Kew Bull. 614.
2*^J:
BM;
specimens deposited however, TYPE: Nu)ts Anhipelagu, Pe-
Pallas's at if 1974). "S. Australia:
391
Number Mosyakin
Volume
3
83,
American
North Salsola
1996
&
Good
trel Bay, Isle St. Francis, 8 Feb. 1802 (fl. fr.) ^ith weak apex; at maturity distinctly winged; fruit-
&
Bauer
(lectotype, selected by Bolschanlzev
4-10
j^^ perianth nun diam.
ca.
BM,
"planta dextra").
(1974),
=
Moquin-Tandon, Chonopo-
Salsola kali L. var. temdfolia Chromosome number. 2n 36 (Mulligan,
dearum Monographica Enumeratio: 136. 1840.
&
Crompton,
1961; Bassett 1970).
TYPE "Ad Baltam
southwestern Ukraine:
locality:
[spbahn. "Balkam" -S. M.] (Besser in herb. DC.)" Vernacular names, Russian-thistle (English);
P Bes-
(holotype, probably at not seen; isotype in the
soude roulantc (French).
KW).
ser memorial collection at
Fenzl Fedebour,
Salsola kali L. var. angustifolia in FI. Habitats, Waste places, roadsides, cultivated
TYPE:
Ross. 798. 1851. no reference to the
3, 2:
disturbed natural and semi-natural plant
fields,
specimens annolated
type the protologne; original
in
communities coastal and riparian sands, semi-
(e.g.,
by Fenzl LE, lectotype not selected.
at
0-2500
and eroded m.
Salsola kali L. var. leptophyila Bentham, Fl. Austral. 5: deserts deserts, slopes);
TYPE
"Queensland and
207. 1870. locality: N. S.
Herbarium specimens were
Distribution.
TYPE:
Wales." not designated.
& and
Salsola tragus subsp. iberica Sennen Pau, Bull. Acad. amined from the foUuvving states provinces:
Intern. Geogr. Bot. (Le Mans), ser. 3, 18: 476. 1908. CANADA: Alberta, British Columbia, Manitoba,
&
Salsola iberica (Sennen Pau) Botschant/ev, Bot. ^^^
Nova Ed-
Brunswick, Scotia, Ontario, Prince
cum
Zhurn. (Leningrad) 54: 991. 1969, aucl. ''Sen-
Q
(Sennen
nen Pau'' (comb, invalid.). Salsola iberica
el
& Pau) Bolschantzev ex Czerepanuv, Svod Dopol- bama, Arizona, Arkansas, California, Colorado,
cum
neniy Izmeneniy k "Flore SSSR": 192. 1973, Connecticut, Delaware, Georgia, Idaho, Illinois, In-
i
aucl. "Sennen et Pau." TYPE: "Hab. Caslille: Mir- Maine,
Kansas, Kentucky, Louisiana,
^^^^^^ j^^^^
anda de Ebro Logrono, terrains vagues, pres
(Elias); ,, «* »«•
.^
Maryliandi, Massachiusetts, Michi i•gan, Maiik-nnesot*a,
\"/ui?
,.. i.
/c
1 station (Sennen) (holotype, not. seen; isotype, iUi^<=):\. J ^ '
la
j
Montana, Nebi raski a, Ntvtevada,
Mississippi, Missouri,
Beck Reichcnbach,
pseudotragus G.
Salsola kali L. var. in
Icon. Germ. Helv. 25: 172. 1909. TYPE: not des- New Hampshire, New Jersey, New Mexico, New
Fl.
ignated; described from Germany, "inprimis in terris York, North Carolina, North Dakota, Ohio, Okla-
interioribus."
Rhode South
homa, Oregon, Pennsylvania, Island,
New
Manual
Salsola pestifer A. Nelson in Coulter's Bot.
Carolina, South Dakota, Tennessee, Texas, Utah,
cVntr?''£ckVMou^^^^^^
Vermont, Washington, West Virginia, Wis-
"The now widely distributed Russian Hustle" (no Virginia,
specimens number
collection cited in protologue; authentic consin, Wyoming. Judging from the limited
GH,
NY, US; lectotype not selected).
at herbarium specimens from Alabama,
available
of
Sornye Ras-
Salsola rulhenica Iljin in B. A. Keller et ah,
Georgia, Louisiana, and Mississippi, this species
teniya SSSR (Weeds of the USSR), 2: 137. 1934,
seems be rare undercollected) in the south-
So6 to (or
nom. Salsola kali L. subsp. ruthenica in
illegit.
So6 Jdvorka, Magyar Nov. Kez. 786. 1951. eastern part of the United States. Salsola tragus
et 2:
TYPE:
not designated. Newfoundland,
probably also occurs in the District
Salsola kali L. var. austroafricana P. Aellen, Mitleilungen
Columbia known from the adjacent part of
of
is
(it
Munchen
der Botanischen Staatssammlung 4: 27.
Maryland), and Florida; native to inland arid
TYPE:
1961. Sudwestafrika: Distr. Liiderit/.-Siid,
Europe and
weed dry riverbed near farmhouse Tarm "Klein regions of southeastern Central Asia,
in
2297 {M
Aus," 28.6.1949, Kinges not seen). follow ^nd occasionally occurs as an introduced alien in
|1
Kew
the synonymy of Botschantzev, Bull. 29: 614.
^^^^^^ regions of the world (naturalized in
^^y^^^
^* South Africa, Australia, and South and Central
Annual herbs, (5-)10-100 cm, glabrous or America),
may have been introduced
sparsely papillose-hispid. Stems profusely ])ranched Salsola tragus first to
from the base or near the base (rarely simple in the United States in South Dakota in 1873 or 1874
underdeveloped specimens), erect, rarely ascend- in flax seed imported from Russia (Piper, 1898;
&
Crompton This
branches normally arcuate. Leaves alternate, Beatley, 1973; Bassett, 1985).
ing;
filiform or narrowly linear, in herbarium specimens noxious weed now^ occupies almost all its potential
mm
range North America. However, young plants are
normally less than 1 broad, semit<'rcte, not in
swollen at base; apex rather soft, subspinescent. In- considered to be an additional forage source for
interru livestock in arid rangelands (Welsh, 1984).
may
lower Bracts maturity reflexed, not im- The mature plant break offal the stem hi
in part). at
±
mu- tumbleweed RoUy-polly
bricate; alternate, abruptly narrowing into to form a (also called in
cronulate-spinose apex. Bracteoles free, or occa- Australia).
The synonymy complicated, and
sionally connate at base in lower flowers. Flowers of S. tragus is
^ome names
applied taxon
solitary or ly 2 or 3 (in the last case lateral the correct use of to this
For example, Botschantzev
uncertain.
flowers mostly abortive) in axils of bracts or is still
who
duced upper leaves. Perianth segments glabrous, (1974), had selected the lectotype of 5. aus-
392
Annals
the
of
Garden
Missouri Botanical
Brown
trails R. (di^posited in BM), t'onsidercd to 3. Salsola paulsetiii D. Litvinov, Turkes-
it I. Izv.
be nonspecific with 5. pestifer and S. ruthenica. tansk. Otd. Russk. Geogr. 01)slich. 28.
4, 5:
Crompton comm.; TYPE: Dominium
(pers. sec also note in Clemants, 1905. 'Turkcstania. Bu-
who
Imd
1992), also studied the type, claimed to chara. In arenosis subsalsis pr. Farab (ad
it fl.
he
conspecific with S, kali sensu stricto. have also Amu-darja), 14 Sept. 1903. Legit Andros-
I A^.
MO.
studied the of aaslralis sent on loan sow M. Koloiv^
typ<' 5. to et (LF])-
herbarium
Tlie she(M contains several fragments.
\x\
Sdlsola peUucidd Lilvinov, Herb. Boss. 49: No
my 1). I. Fl.
opinion, four of fragments belong
tliese to S,
T^
2434. 1913. IMv "Dominiuin
arenosis
liiicliara, in
kali subsp. pontira, a southern maritime race of the
1913—
pr. Farab. 14 Sept. 1903 [sphahn. S. M.].
S. kali aggregate (see discussion above). However, Leg. Androssow''' (LF).
A^.
the lectotype of australis selected Botschantz-
.S, l)y
Annual 10-80(-100) cm,
herbs, glabrous or
ev (1974) the fragment, which mounted
is 111th is
sparsely papillose-hispid. Stems profusely
I.
-J »u ^M
at, t,1he ri• ght side otf the 1herIbar•ium sIheet*. Ihi•s Irrag- /
,'
r
brancliiedi from tihe bi ase or near the bi ase /
(rarely
ment
represents a form morphologic-ally interme-
simple underdeveloped specimens),
in erect, rarely
diate between 5. kali su\)^p. ponlica and tragus.
.S.
ascending or prostrate; branches straight or arcuate,
The
combination
kali subsp. tragus
5. (L.)
almost Leaves
oftiMi p<*ri)(*ndicular to the st(»rn. al-
Celakovsky was
constantly misap[)lied Europt*
in lo
ternate, filiform or linear, in herbarium specimens
the littoral taxon treated here as S. kali subsp. pon- mm
normally ca. broad, but occasionally slightly
1
discussion under
tica (see that laxon). Th(* author-
broader, semitercte, not swollen base; apex spi-
at
ship of the former combination was incor-
oft<*ii
nose. Inflorescence splcate, distinctly interrupted
at
rectly attributed Nyrnan (Consp. Europ.: 631.
to Fl.
maturity. Bracts at maturity strongly reflexed, not
who
1881), in fact validated 10 years after
„i
it •
imhn..c- at,e;. ali,ternat»e, iiaiTowi• ng •nit, o spinose apex.
x^
^
eiaKovsKy.
Bradeoles free or connate near \hv base, spreading,
In North America, taxon was subsequently
this
spinescent. Flowers solitary, or rarely 2 or 3 (in the
treated as kali var. tenuifolia (with the authorship
.S. l-j^t case lateral flowers mostly abortive) in axils of
who
incorrectly attributed Tausc^h, published
to
\n-iw\^ or reduced upp(T leaves. Perianth segments
only the name, nomen nudum),
.S. prstifer, S. iberica glabrous, with long-acuminate spinose apex, ma-
at
(with authorship incorrectly attributed Senncn
its to forming slender columnar beak above
turity a the
&
who
Pau, publislnMl their taxon as a sulispecies), 7-12
wings, prominently wingecL Fruiting perianth
and S. australis. In Europe this taxon was also m am.
di
known
and
as ruthenica kali subsp. ruthenica
5. S.
—
Chromosome
number. 2n 36 (Semiotrocheva,
Soo. Salsota ruthenica an illegitimate
(Iljiii) is
1983; see
fig. 3).
name, when
because, describing his species,
Iljin
synonymy
cited in the earlier valid
its S, pestifer,
Vernacular name, Barbwire
Russian-thistle.
name
The name
of the sanu^ rank. subspccific pro-
posed by Soo basionym Habitats. In sandy soil in disturbed natural and
is legitimate, but, sinc(^ its
open
^t'"ii-"atural plant (H)mmunities sands,
was new (e.g.,
be
illegitimate, siiould regardiul as a
it
and
name new ICBN ^^^"^' *^""^^' ^^"*'y ^^^^^^ places, semi-deserts
(not a combination; see article 58,
0-1900
deserts, eroded sandy slopes, m.
etc.);
Greuter
et 1994).
al.,
Recently Tzvelev (1993) confirmed that the cor-
Herbarium specimens were
Distribution, ex-
name
rect for the widespread narrow-leaved weedy
amined
from
the following Arizona,
Califor-
stat(*s:
representative of the kali aggregate tragus
S. is
.S. nia, Colorado, Nevada, Utah. Salsola paulsenii
is
L, name
a that had already been accepted for this
„^,ive to soutbeasternmost Europe and Central
taxon by Degen (1937). Judging from the photo-
Asia.
graphs of Limiaean specimen of tragus
the* S. This species confined mostly open sands,
to
is
(LINN
which
315.3), selected as the lectotype,
may
I .,nd rarely to saline sandy habitats. be ex-
It
this point of view correct. This cliange of a nanu New
is [)ected in the future in Mexico and Texas, as
seems to be desirable, because would guarantee some
it well as in Great Plains states. was
It first
more
nomenclature
stable for this taxon in the fu- reported from North America by Muiiz and
(1968),
ture. The name tragus was used for this species the determination was made by Ad-
.S. Biitschantzev.
by some American botanists in the 19th century, and
ditional details of distribution mor{)hology of
but unfortunately, most North American botanists this species have been discussed by Beatley (1973)
common
did not resist the temptation to accept the and Fuller
(1986).
European
misapplication of the nanu\ Salsola paulsenii weakly differentiated from
is its
Volume Number Mosyakin 393
3
83,
1996 American
North Salsola
allied taxon, 5. tragus sensu stricto. The interme- fied. Later spread to Colorado, Iowa, and Missouri
it
diate forms between them seem be quite rare in (Cory, 1948; Schapaugh, 1958; Muhlenbach, 1979).
to
common
Central Asia, but more southeastern- At present species known North Amer-
in this in
is
most Europe, western Kazakhstan, secondary syn- ica mostly from the Great Plains region and scat-
anthropic localities in East Europe, as well as in tered localities in other states. Recently was also
it
&
the United States. discovered in Canada (Crompttm Bassett, 1985).
However, seems be under-
actual distribution to
its
common
4. Salsola collina P. S. Pallas, lUustr. PL: 34. estimated due to the and constant confu-
TYPE:
1803. "in tractu collium cotaceorum sion with deviate forms of tragus, w^hich occa-
S.
Rhymnum
Samaram
inter et a jugo Uralensi sionally resemble S. collina in having narrow
fl.
BM
descendentium'' (holotype, not seen). inflorescences and gall-like caducous flowers/fruits
at the axils of lower and middle branches. In the
Annual herbs, 10-100 cm, sparsely densely
to
may
future S. collina be expected to occur within
papillose or hispid (rarely almost glabrous). Stems
the present range of tragus throughout North
S.
branched above the base (occasionally with slender
A
menca.
branches near the base), erect, rarely ascending;
was also reported as a casual alien from sev-
It
branches Leaves
straight or slightly arcuate. alter-
eral countries of western and central Europe and
is
narrowly some-
nate, filiform or linear, semiterete,
regarded to be established or even completely nat-
times semi-amplexicaul base; apex with rather
at
map
Europe second-
uralized in eastern (see of
its
soft bristle (rarely spinescent). Inflorescence nar-
&
ary distribution in Baranova Khilova, 1990),
rowly spicate, not interrupted. Bracts at maturity
appressed and imbricate, alternate, gradually nar-
soda PL
5. Salsola Sp. 233. 1753. Kali
L., 1:
rowing into a subulate spinose apex. Flow^ers soli-
soda Scopoli, FL Cam., ed. 175.
(L.) 2, 1:
tary or rarely 2 or 3 in axils of bracts or reduced
TYPE: LINN
1772. 315.7 (lectotype, selected
upper leaves (sometimes flowers are also present
at
Hedge
by I. in Jarvis et al., 1993). [The spe-
axils of lower leaves and branches, at maturity
cies described from southern Europe; "Habitat
forming gall-like caducous balls). Perianth seg-
Europae
in australis salsis."]
ments glabrous, with weak, flaccid apex; maturity
at
wingless or with narrow erose wing, becoming con- Annual herbs, 5—70 cm, glabrous. Stems
nate and united with bracteole bases; fruiting peri- branched from the base or nearly so, erect or as-
mm
anth ca. 3-5(7) diam. cending; branches straight or slightly arcuate (lower
ones sometimes almost prostrate). Leaves (especial-
= &
Chromosome number. 2n 18 (Pohl Gilles-
ly lower ones) mostly opposite, linear, semiterete,
pie, 1959).
herbarium specimens more
usually
fleshy, in
tl
mm
Habitats, Waste places, roadsides and railway 1.5 broad, distinctly swollen or ovate base;
at
areas, cultivated fields, disturbed natural and semi- apex mucronulate, non-spinose. Inflorescence spi-
100—2000
natural plant communities; m. cate, distinctly interrupted. Bracts at maturity hor-
(?)
izontally reflexed, alternate or almost opposite,
Herbarium specimens were
Distribution. ex-
abruptly narrowing mucronulate non-spinose
into
amined
from the following states in the U.S.A.: Ar-
apex. Flowers mostly solitary in axils of bracts or
izona, Colorado, Iowa, Kansas, Michigan, Minne-
reduced upper leaves. Perianth segments glabrous,
New
Missouri, Montana, Nebraska, Mexico,
sota,
with crenate to pectinate-ciliate apex, at maturity
North Dakota, Texas, Utah, and Vermont.
wingless or with rudimental triangular tubercles,
The
species also reported from additional
is lo-
3—
not connate to bracteoles. Fruiting periantli ca.
Iowa (Schapaugh, Utah
calities in 1958), L.
(S. mm
diam.
6(7)
Welsh, comm.), and Canada: Ontario
pers. (possibly
&
also in Quebec) and Saskatchewan (Crompton Chromosome number, 2n 18 (Zakhar'yeva,
Bassett, 1985). Salsola collina native south- 1985).
is to
easternmost Europe, southern Siberia, and arid
and
Habitats, Coastal disturbed saline habitats;
known
regions of Central Asia; as a naturalized
is
it
0-50
m.
or casual alien in some other regions of Europe and
Asia. Distribution, Herbarium specimens were ex-
This Asian species was reported for the time amined from California: San Mateo Co., Palo Alto
first
for North America from Minnesota by Moore Yacht Harbor near airport, altitude about sea level,
(1938). However, had been collected in Kansas 6 Oct. 1974, H. Thomas 17615 (MO); Santa Cla-
it J.
in 1923 (Brooks et al., 1976), but was misidenti- ra Co., Palo Alto Yacht Harl)or, near E end of yacht
394
Annals
the
of
Garden
Missouri Botanical
edge
basin, of dislurhed area along salt marsh, 7 tential forage plant introduced from Syria in 1969.
Dec. 1975, Thomas 18062 (US). Sahola soda Salsola vermiculala designated as a Federal Nox-
/. //. is
Weed
is native to Eurasia (mostly Atlantic Europe, the lous l)y the Unit<Ml States Department of Ag-
Mediterranean region, and southwestern Asia) and rieulture (Westbrooks, 1993). so far the only
It is
northern Africa. perennial s])eeies of Salsola sensu lato introduced
known
Salsola soda is from several localities in to North America. Together with other related Eur-
central California, near San Francisco Bay (Tho- asian an<l African laxa, should probably be seg-
it
as, 1975). can be expected to spread in Cali- regated into a separate genus, Caroxylon.
It
foniia, or to app(*ar in inland or coastal saline hab- Salsola lermiculata sensu lato a taxonomlcally
is
itats in other southern stat<*s. complicated and morphologically polymorphic
complex Med-
represented in Eurasia (mostly in the
6. Salsola veriniculata L., Sp. PL: 223. 1753. iterranean n^gion, western and Central Asia) by
TYPE: LINN
315.20
(leetotype, selected by several closely related races usually treated as sub-
Botschantzev, 1975b). [The species described species or distinct species (R(»tschantzev, 1975a, b;
from Spain: "Hispania.''] Greut(M- et 1984). S(dsola vermiculata sensu
al.,
stricto a w<^stern Mediterranean species occur-
is
20—
Subshrubs, 70(— cm, densely pubescent
100)
southwestern Europe ami
ring in northwest<*rn Af-
(especially wlien young) with smooth and minutely
Numerous
records of vermiculata from Syria
rica.
.S.
denticulate hairs; sometimes becoming glabrous
at
and oth(M* countries of the Middle East refer to other
branched woody
maturity. St(Mns the base,
at
closely related species and/or subspecies, which
re-
5—8
branches erect or ascending, virgate. Leaves
Meditenanean
place in the eastern region. North
it
X mm,
0.5-1
semiterete, usually pubescent, ex-
Anu^rican most probably belongs da~
mal(^rial to 5.
panding
Into ovate base, bearing in their axils sev-
mascena sensu the protologuc and the
stricto.
1^ mm It fits
eral reduced leaves ca. long; apex ol)tuse.
type spi'cimen depositerl LE, but additional
at
Inflorescence spicate but primary axis some-
its
study and comparison with other Eurasian "micro-
limes pani(*ulately branched. Bracts normally
Some
species^' are necessary. of these taxa remain
densely pubescent, obtuse. Bracteoles Flowers
free.
little-known and poorly understood taxonomically.
2-3
reduced
solitary or rarely in axils oi bracts or
Because
of that prefer provisionally to place the
I
upper leaves. Perianth segments sparsely pubes-
North American plant of Syrian origin in S, ver-
cent above wings (especially apex), sometimes
at
miculata sensu
lato until further clarification of
its
becoming winged
glabrous, with conical apex,
at
taxonomic
ich'utity.
7—12
maturity. Fruiting perianth (iticluding wings)
The
damas-
nonuMiclatural citation for Salsola
mm
diam.
and synonymy
cena basic are provided below.
Its
=
Chromosome
number, 2n 18 (Sankary, 1986;
damascemi
Salsola Botschantzev, Bot. Zliurn. (Len-
th e recordf.or 5. vermiculata L. van lillosa (Del.)
TYPE:
ingrad) 500. 1975. Tallus
Moq."). 60(4): ^^Syrie:
pien'eux du jardin Boustan Nashe a Mezze
el
Rocky
Hahitats, slopes, clay disturlxnl
soils,
pr^s dc Damas, 9 Aug. 1856, GaiUardot
C.
1000
places; ca.
ni. 162r
(holotype, LE).
Herbarium specimens were
Distribution, ex-
Acconling Botschantzev (1975a. the syn-
to b),
amined from California: San Luis Obispo Co., Re-
damascena
^^„y„,y of Salsola includes rigida Pal-
5.
W
km
Grade Kern Tem-
cniit Pass, of Co. Hne,
5^2
\^^ ^^^ tenuifolia BoissicM* (Flora Orient. 4, 2: 968.
L
979
blor Range, elev. m, 3 Oct. 1978, Johnson
/. and
1879, vermiculata subsp.
in [)ait) S, tenuifolia
&
G. D. Rarbe 2448 (US); Temblor Range, ridge
(Boissier) Bots<'hantzev (Novosti Vyssh. Rast.
Sist.
E
top and slope Crocker Grade, S of Crocker
at
The names
(Leningrad) 375. 1964). Salsola
1: vil-
W
Springs Rd.. ca. 2 mi. of county bne and 7.5
tlu^
/^,.^^ d^.j;]^. (p] Aegyi>t. llluslr.: 57, No. 309. 1813,
SW
3200
mi. of McKittrick, 31
(airline) <dev.
ft., in part, excluding the and vermiculata
typ<^) 5.
Don
Oct. 1989, Pendleton (MO). Salsola
s,n. ver-
subsp. Eig sen
villosa (Delile) (Palest. Bot.,
3,
J. J,
miculata native the Meihterranean region.
is to
3: 132. 1945, in part, excluding type of the
tlie
known
Salsola vermiculata sensu North
lato In
is
basionym) were misapplied damascena sensu
to S.
America only from Califonu'a, as a locally persis-
stricto.
tent escaped weed. naturalized near an aban-
It is
doned experimental Recruit Grade Pass, T(Mn-
plot. Cilrd
Literaliirr
Range, San Obispo
blor Luis Co. (possibly also in &
Baratiova, E, V. E. V. Khilova. 1990. Materialy k geo-
Kern when^ was
Co.), previously tested as a po- graficlicskoy kharakterislikt' lU'kutorykh pn*(lstaviteley
It
Volume Number 3 Mosyakin 395
83,
1996 American
North Salsola
&
antropofil'nogo eleiueiita flory Severo-Zapada evropeys- Jafri, S. M. H. V. B. Rateeb. 1978. Chenopodiaceae
SSSR 1-109
koy chasti (Materials for geographical charaeter- Pp. in Flora of Libya. Vol. 58. Al Faateh Univ.,
some
istics of representatives of the anlhropophiloiis flo- Fac. Sci., Dept. Rot., Tripoli,
&
ristie element in the Southwest of the European pari of Jarvis, C. E., F. R. Rarrie, D. M. Allan L. Reveal.
J.
A
the USSR). Vestn. Leningradsk. Univ., Ser. 3 (Riol.), 1993. list of Linnaean generic names and their types.
Regnum
35-44. 1-100.
Veg. 127:
1(3):
& &
Bassett, C. W. Crompton. 1970. In: A. Love (ed- Jonsell, B. C. E. Jarvis. 1994. Lectotypification of Lin-
I. J.
lOPB chromosome number XXVll. Taxon naean names Nordica (Lycopodiaceae
reports. for Flora Vol.
itor), 1
437^42.
Papaveraceae). Nordic 145-164.
19: Bot. 14:
J.
Beatley, C. 1973. Russian-thistle (Salsola) species in Moore, W. 1938. Euphorbia dentata Salsola collina
J. J. ixinl
225-
western United Range Managem. 26: Minnesota. Rhotlora 40: 135.
Slates. in
J.
226. Muhlenbach, V. 1979. Contributions to the synanthropic
R
Rod
Botschantzev (Bochantsev), V. 1969. Salsola L., (atlventive) flora f>f the railroads in St. Louis, Missouri,
kratkava istoriya ego razvitiya rasseleniya (The genus U.S.A. Ann. Missouri Bol. Card. 66: 1-108.
i
A
Salsola concise history of development and Mulligan, G. A. 1961. Chromosome numbers of Canadian
L.: its
dispersal). Rot, Zhurn. (Leningrad) 54(7): 989-1001. weeds. III. Canad. Bot. 39: 1057-1066.
J.
A
1974. synopsis of Salsola (Chenopodiaceae) Munz, A. 1968. Supplement to a California Flora.
P.
.
Kew 597-
from South and South-West Africa. Bull. 29: Univ. California Press, Berkeley.
614. Pallas, 1803. Illustrationes plantarum imperfecte vel
P. S.
1975a. Novye vidy Salsola (New species of nondum cognitarum, cum centuria Iconum. De Halo-
L.
.
Salsola L.). Bot. Zhurn. (Leningrad) 60(4): 498-505. phytis, seu plantis apetalis kalicis generalim. Lipsiae.
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& P
cies subsectionis Vermiculatae Bittsch. sectionis Carox- Pohl, R. W. Gillespie. 1959. Distributional and
J.
265—
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160-194.
Vyssh. Rast. 12: 267.
&
An Chromosome number
Britton, N. L. A. Brown. 1913. Illustrated Flora of Sankary, M. N. 1986. reports 91.
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New new
Possessions. Ed. 2. York. Schapaugh, W. 1958. Salsola collina Pall., to Iowa.
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Brooks, R. E., R. L. McGregor L. A. Hauser. 1976. Proc. Iowa Acad. Sci. 65: 118-121.
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Vascular plants to the state of Kansas. Pp. Semiotrocheva (Semiotroczeva), N. L. 1983. Kariosiste-
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in: Records of the Fauna and Flora of Kansas for matika nekotorykh \idov rodov Salsola L. Climacop-
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