TAXONOMY OF CUSCUTA GRONOVII AND CUSCUTA UMBROSA (CONVOLVULACEAE)

2006·7.1 MB·English

by  Mihai Costea

TAXONOMY AND
OF CUSCUTA GRONOVII
UMBROSA
CUSCUTA (CONVOLVULACEAE)
Guy Nesom
Mihai Costea
(corresponding author)
L.
Deportment of Biology Botanical Research ofTexas
Institute
509 Pecan
Wilfrid Laurier University Street
75 University Avenue West Fort WorthJexas 761 02-4060, U.S.A.
Waterloo, Ontario [email protected]
N2L3C5XANADA
[email protected]
Sasa Stefanovic
Department
of Biology
University of Toronto at Mississouga
3359 Mississauga Road
Mississouga, Ontario
L5L1C6XANADA
ABSTRACT
C
name
Cuscuta umhrosa the correct for the species often recognized as megalocarpa. appar-
is It
ently intergrades in some features with C.gronovii, but the two are generally distinct across a broad
region of sympatry. Three varieties are recognized within C. gwnovii: \3.r. gronovii is widespread; var.
broadly sympatric with gronovii but maintained because relationships with
laliflora is var. is C.
C
gronovii var. gronovii and cephalanthi are not clear; var. calyplrata is more distinct and narrowly
although sympatric with
restricted in range, also var. gronovii.
it is
RESUMEN
Cuscuta umbrosa es el nombrc correcto para la especie reconocida recientemente como C. mcgahcarpa.
Es ampliamente simpatrica y algo intergradada con C.gronovii. pero las dos especies son generalmente
muy
claras. Se reconocen tres variedades dentro de C. gronovii: var. gronovii que esta esparcida; var.
C
laliflora es ampliamente simpatrica con la var. gronovii pero se mantiene debido a la relacion con
mas
no estrechamente
gronovii var. gronovii y C. cephalanthi esta clara;la var. calyplrata es clara y
restringida en su distribucion geografica. aunque sea tambien simpatrica con la var. gronovii.
C
The original concept of Cuscuta umhrosa Beyr. ex Hook. (1840) included
&
gronovii Willd. ex Roem. Schult. (1820) in typical form as well as a variant
Engelmann
plant, based on the specimens cited in the protologue. (1859) clari-
C
from and
fied the typification of umhrosa, distinguishing C. gronovii, de-
it
scribed the variant as C. gronovii var. curta Engelm. Yuncker recognized the
name
variant taxon at specific rank, at first (1921, 1932) using the C. curta
C
(Engelm.) Rydb. but later (1943, 1965) adopting umhrosa as the earliest legiti-
C
We
mate name. find that recently umhrosa either has been accepted as a spe-
cies under yet another name, C. megalocarpa Rydb. (1901) (Kartesz 1999;
SIDA22(1):197-207.2006
198 BRIT.ORG/SIDA 22(1)
NRCS
Darbyshire 2003; USDA, 2004), or else has not been recognized at any
it
&
taxonomic rank (Beliz 1986-1987, in herb; Crins Ford 1988).
C
Cuscuta gronovii and umhrosa, as recognized here, are similar and ap-
C
parently closely related to ccphalanthi Engelm. and rostrata Shuttlw. ex
C.
&
Engelm. A. Gray. These four species, which were placed by Yuncker (1965) in
two subsections (P/at3/carp67e and Cepha/cinthae), have the same pattern of stem
growth and branching: haustoria are formed as the main stem twines around
the host and no tendril-like axillary branches are generated. This pattern was
m
Dawson
observed by (1984) C.^ronovii. The evolutionary and taxonomic
first
C
status of umhrosa reevaluated here, especially with regard to relation-
is its
with
ship C.gronovi'L
METHODS
Descriptions of morphology are based on samples from specimens from DAO,
SMU
NY, and (Appendix Measurements and pictures were taken with scan-
a
1).
KV
ning electron microscope Hitachi S-570 Samples were coated with 30
at 15
nm
K
gold using an Emitech 550 sputter coater. Terminology regarding the mi-
cromorphology and and
of flowers, seeds capsules, pollen were described in
detail in the paper published in this issue (Costea 2006). Conserva-
first et
al.
was
tion status assessed using NatureServe (2005) ranks and
criteria.
TAXONOMY
Yuncker
(1932, 1965) characterized Cuscuta gronovii as having "styles about
somewhat
equaling the ovary or shorter," in contrast to those of C. umhrosa,
which
are "very short, about one fourth the length of the ovary" Other con-
trasts between these taxa emphasized by Yuncker are in the morphology of
C
infrastaminal scales and the size of capsules and seeds. According to him,
umhrosa has infrastaminal scales about half the length of the corolla tube, broad,
and and and
often truncate biiid, capsules seeds are larger. In contrast, C.gronovi
i
has infrastaminal scales about equaling the corolla tube, oblong and not trun-
We C
and and
cate or bifid, capsules seeds are comparatively smaller. find that
C
gromwii and umhrosa
usually are morphologically distinguishable, but
C.
C
may show when
gronovii character states similar to umhrosa even occurs
it
C
outside the geographical range of the For example, gronovii from On-
latter.
may
tario and Quebec occasionally have more or truncate infrastaminal
less
scales one-third the length of the corolla tube, or short stigmas, or comparably
&
Commonly
large seeds (Crins Ford 1988) and capsules. only one char-
state
um
acter o[ C. hrosa, or rarely a combination of two large capsules and seeds),
(e.g.,
Such may
occurs in gronovii plants of C.gronovi be separated from umhrosa
C. C.
i
using the character combinations in the following key. Other differences can
be found in pollen morphology: gronovii has pollen grains with tectum
C.
COSTEA ET AL. JAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 199
C
imperforatum few puncta, while umhrosa has tectum perforatum
or a or
al-
most
microreticulate
(Fig.
1).
Infrastaminal scales oblong,apex usually rounded, not bifid, rarely truncate, ca. reach-
1.
mm
ing the filannent bases, rarely ca. 1/3 of the corolla tube; styles (0.6-)1.2-2.2
X mm; X
long; capsules 2.5-4.5(-5.2) 2-4(-5) seeds 135-1.7(-2.4) 1.2-1.5(-1.6)
I.e. gronovii
Infrastaminal scales broadly-oblong, apex truncate to slightly bilobed, l/2(-1/3) as
1.
mm
X
3-5
long as the corolla tube; styles 0.3-0.7(-0.9) long; capsules 3.5-6. 5(-7)
mm
X
2.C.umbrosa
mm;seeds(1.8-)2-2.5(-2.8) 1.5-1.65
(-6)
Cuscuta gronovii has a wide geographic distribution in North America and
is
C
common
broadly sympatric with umhrosa. The former more in central and
is
northeastern regions (Yuncker 1932), while the latter primarily a prairie species.
is
C
Cuscuta gronovii usually distinct from umhrosa by a greater morphological
is
C
among
ap than are the varieties of gronovii themselves (see key below).
Within Cuscuta gronovii, Yuncker (1965) recognized three varieties: van
gronovii, var. latiflora Engelm., and var. calyptrata Engelm. Variety latiflora and
show
variety gronovii are broadly sympatric across their geographic range, they
no distinct preferences toward hosts, and morphological intermediates occur.
Var. calyptrata more narrowly restricted in range (Texas and Louisiana) but
is
sympatric with the widespread gronovi Continued recognition of sym-
is var.
i.
unusual
patric varieties within Cuscuta gronovii reflects the observations
is It
among
morphological each
that a close similarity exists the varieties, (2) of
(1)
may
the varieties usually morphologically distinct although intermediates
is
be relatively common, and each of the non-typical varieties shows features
(3)
suggesting that might be more closely related to species of the Cuscuta
it
among
cephalanthi-rostrata group or at least that genetic interchange taxa of
C
may
broader group have influenced the variation patterns within
this
We
gronovii. consider premature to discard the varieties as formally recognized
it
taxonomic entities until relationships involving all these taxa are clarified.
CuscutagronoviiWilld.exRoem.&Schult.,Syst.Veg. 1820
1. 6:205. (Fig. 1 a, c,
Roem.&Schult NieuwL&Lunell^Amer.
f). Epit h_ym u m gronovi i(W'i lid. ex J Mid). Naturalist
4:511. 1916. Typi^ (see Yuncker 1943); North America, unspecified locality, Willdenow 3160-10
photo
(B, a seen).
medium
Stems to coarse, yellow to orange. Inflorescence of pedicellate (1-4.5
mm)
cymose sometimes endog-
flowers in loose or dense, paniculate clusters,
enously formed; each with one ovate broadly triangular bract the
low^er to at
f
pedicel base, and sometimes l-2(-4) bracteoles on the pedicels. Flowers 5-merous
mm
(sometimes 3- or 4-merous), 2-4(-4.7) long, not papillose, white-cream,
many
with a few to isolated, round to ovoid translucent laticifers in the calyx,
and Calyx campanulate narrow-campanulate, reaching
corolla, ovary. cupulate, to
middle divided 1/2-2/3
to ca. the of the corolla tube or the corolla sinuses, ca.
200 BRiT.ORG/SIDA
22(1)
Fic. 1.a-b. Capsules with calyx at the base: a. Cuscuta gronoviiydir,gronovii; b. Cuscuta umbrosa (assembled from two
=
=
pictures) (scale bar mm); c-e. Morphology of pollen. c.fuscufogra/joW/ (scale bar 7.5 jam); d-e,fy5cwf£7wm6roso
1
= 8i =
(scale bar arid 3 j.im,respectively);f-h.Morphologyofseeds:f.C.^ronow7;g.f.™ftro5ff (scale bar 0.6 mm); h.
=
C
Hydrated seed coat of umbrosa (scale bar 120
)J.m).
COSTEA ETAL., TAXONOMY OF CUSCUTA GRONOVII AND UMBROSA
C. 201
the calyx length, lobes ovate to suborbicular or oblong, apex rounded or ob-
more
tuse, or less basally overlapping, margins entire to serrulate; corolla tube
mm
campanulate, broadly- narrow campanulate
or
(l-)1.5-2.5(-3) long, lobes
commonly
mostly
ovate, rounded-obtuse, 1/3 the tube corolla tube, sometimes
wax
1/2 to equaling the corolla tube, spreading or ref lexed; epicuticular absent
or reduced to a few reticulated rodlets; stamens exserted, anthers on filaments
equaling or longer than anthers; pollen 3(-4)-zonocolpate, (18-)19-24(-27) jim
rounded truncated polymorphic same
long, or at poles, in the anther or flower,
from spheroidal subprolate former shape more tectum
to (the frequent);
imperfortatum with a few puncta, rarely tectum perforatum with puncta 03-
0,5 )im in diameter; sexine scabrate usually with isolated granules; infrastaminal
scales oblong or broadly-oblong, rounded or truncate and deeply fringed
bifid,
distally, shorter than the corolla tube or reaching the filament bases; styles dis-
tinct, slender, thickened at the base or occasionally slightly subulate, (0.6-)1.2-
mm
2.2 long; stigmas capitate, globose. Capsules globose, ovoid globose-conic
to
or subobpyriform, thickened at the top, interstylar opening relatively small,
sometimes with a short and broad but discernible neck, 2.5-4.5(-5.2) x 2-4(-5)
mm,
indehiscent or irregularly dehiscent, surrounded or capped by the with-
2-4 x
ered Seeds per
corolla. capsule, 1.35-L7(-2.4) dorsoventrally
1.2-L5(-1.6X
compressed to obscurely angled, subrotund to broadly-ovate; hilum subtermi-
x mm,
nal, subrotund, 0.33-0.39 0.30-0.33 vascular scar of funiculum
linear,
mm
0.20-0.25
long, vertical to slightly oblique. Surface of seed epidermis vari-
when when
able: alveolate dry and papillate hydrated (cells 20-50 jam in di-
more wax
ameter), or cells or less polygonal and puzzle-like, with epicuticular
organized as reticulated rodlets. 2n = 60 (Fogelberg 1938).
KEY TO VARIETIES OF CUSCUTA GRONOVII
Calyx tube about equaling corolla tube; calyx lobes oblong to ovate, barely over-
1.
mm
lapping, corolla tube 1-1.5 long, broadly campanulate, corolla lobes ca. 1/2 to
equaling corolla tube gronovii
b. C. var. latiflora
Calyx ca. 1/2 of corolla tube, calyx lobes ovate to suborbicular, overlapping;corolla
1.
mm
tube campanulate narrow-campanu
1.5-2.5 (-3) long, to late; corolla lobes 1/
3(~l/4) of the corolla tube.
mm
Corolla tube narrow-campanulate, 2-2.5 (-3) long, withered corolla cap-
2.
ping capsule gronovii calyptrata
C.
c. var.
mm
Corolla tube campanulate, 1.5-2.5 long, withered corolla surrounding the
2.
capsule base before falling a. C. gronovii var gronovii
Cuscuta
gronovii
gronovii.
a. var, Cuscuta vulgivaga Engelm., Amer.J. Arts 43:338. 1842.
Sci.
&
Cuscutagronovi WiUd. ex Roem. Schult. var. vulgivaga (Engelm.) Engelm., Trans. Acad.
Sci.
i
Engelmann
St. Louis 1:508. 1859. (1842) described three varieties within the species and cited
specimens for each of the three (as below), but he did not specify which variety was typical
for the species nor did he cite any specimens pertinent to the typical expression of the spe-
cies, exclusive of varieties. In 1859, he noted for C.gronovii var. vu vaga that Willdcnow's
Igi "It is
hisHb. 3160"
original C.gronovii, in nro. (thus equating var vulgivaga with var. gronovii).
202 BRIT.ORG/SIDA 22(1)
Lectotype
Cuscuta vulgivaga Engelm. var. [_ iaxifiora Engelm., Amer. Sci. Arts 43:338. 1842.
J,
]
(designated here): U.S.A. Alabama: [Oct] 1824, Buckley s.n. (MO). Engelniann cited three (or
New
York on Decodon, Gray; Missouri on Ccphalanthus
four?) collections: ^'Western Dr. A.
and Amphicarpaca, and Georgia, on "? Carey; Alabama, on Salix and Asicr., S.B.
[sic],
J.
Buckley" The lectotype sheet lias a branch of Salix nig}-a and one of Symphyolrichum sp.,
each hosting Cuscuta.
LECTOTYPE
Cuscuta vulgivdga Engelm, vdTA_\glomcrata Engelm., Amer.J.Sci. Arts 43:338. 1842.
On Engelmann
Vermont: Carey (MO).
(designated here): U.S.A. Leersicu [no date], Johfi s.n.
New
noted that "my specimens are from Vermont on Leersia, and Hampshire, on Solidago,
both from Mr. Carey" The lectotype specimen has a small branch of the host with Cuscuta
J,
and a packet with dissected flowers. Mounted on the same sheer is a fragment packet with a
New
Carey specimen of Cuscuta "on Solidago" from Hampshire. appears likely that both
It
specimens fragments
arc of the original collections.
Cuscuta Engelm. tciramcra Engelm., Amer.J. Arts 43:338. 1842. TYPE: U.S.A:
vulgivtiga var. [_] Sci.
MO,
Engelmann noted only "Connecticut, on Urtica,}. Carey" (holotype; presumably, not
located).
common
and The most and widespread dodder North
Dist hut-ion eco Iag_y.— in
ri
CANADA:
America (Engehnann 1859; Yuncker 1965). Alberta, British Colum-
New
Nova
bia, Manitoba (only a few collections), Ontario, Brunswick, Scotia,
Prince Edward Island, Quebec, Saskatchewan (only a few collections). U.S.A.:
Columbia,
Alabama, Arkansas, Arizona, Colorado, Connecticut, District of
Florida, Georgia, Iowa, Idaho, Indiana, Kansas, Kentucky, Louisiana,
Illinois,
Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri,
New New New
North
Montana, Nebraska., Hampshire, Newjersey, Mexico, York,
Rhode
Oklahoma,
Carolina, North Dakota, Ohio, Oregon, Pennsylvania, Island,
South Carcolma, South Dakota, Tennessee, Texas, Vermont, Virginia, Vvfest Vir-
Wisconsin, Wyoming. West Indies (Yuncker 1965) and Europe along wa-
ginia,
courses (Feinbrun 1972). Growing in a wide variety of climates but appar-
ter
shaded and/or wet environments, sometimes occurring
ently preferring as a
&
weed, in cranberries and grapevine (Parker Riches 1993). Flowering Jun-
e.g.,
Nov Hosts: numerous species from liundreds of genera of herbaceous and wood
plants in various families, including Acanthaceae, Anacardiaceae, Apiaceae,
Commelmaceae,
Asteraceae, Bignoniaceae, Brassicaceae, Caprifoliaceae,
Convolvulaceae, Cornaceae, Balsaminaceae, Betulaceae, Euphorbiaceae,
Fabaceae, Lamiaceae, Polygonaceae, Primulaceae, Rosaceae, Rubiaceae, Solan-
and Gaertner
aceae, Urticaceae, Verbenaceae, Vitaceae, others (see also 1950).
m
G5 (common) same
Conservation assessment NatureServe 2005).
status: (the
b. Cuscuta gronovii var. latiflora Engelm., Trans. Acad. Sci. St. Louis 1;508. 1859.
LEL:roTYPH U.S.A. N'liSSOURl. Margin of lakes and swamps, in the "American Bottom'^ oppo-
():
Saint Louis, on Saururus, Sep 1841, Geyers.n. (MO, designated by Yuncker 1921), Engel-
site
m
mann synonymy
cited "Csau/'tiri, Engelm. r ol Cuscuta gronovii var. latiflora.
&
Rocm. Schuk.
Cuscuta rini Engelm., Atner. Sci. Arts 43:339. 1842. Ciiscutagronovii Willd. ex
S(7!i J.
saururi (Engelm.) MacMill, Metasp. Mmnesota Valley 430. 1892. Llctotypf. (Yuncker
var.
margm
1932J: U.S.A. MISSOURI: of lakes and swamps, in the "American Borrom^' opposite Saint
COSTEA ET AL. JAXONOMY OF CUSCUTA GRONOVM AND C. UMBROSA 203
Louts, on Saururus^ Sep 1841, Geyer s.n. (MO). Engelmann also cited ''Alabama, Dr. A. Prout;
Lmdheimer"
Texas, on Boehmeria, Polygonum, &€-, F
and intermediates with gronovii are relatively
Variety variable var.
latijlora
is
common. In seemingly intermediate specimens, on a single plant flowers can
somewhat
be found with the calyx about equaling the corolla tube or shorter.
Apparently the corolla tube may grow to some extent after fertilization occurs,
thus modifying the ratio between calyx length and corolla tube. In var. gronovii
The
the flowers have the calyx about half the corolla tube length. size of
all
C
flowers and capsules in latijlora similar to cephahnthi, but the latter
var. is
4-merous flowers and capsules capped by the withered corollas.
differs in its
m
m
m
mostly depressed-globose capsules with large intersylar apertures.
differs
its
ijl
m
Distribution and ecology—The same distribution and hosts as Yar. gronovii
common.
Flowering Sep-Nov.
but
less
(common) same assessment NatureServe
Conservation status.-G5 (the in
2005).
Cuscuta gronovii var calyptrata Engelm., Trans. Acad. Sci. St. Louis 1:508. 1859.
c.
Cuscuta calyptrata (Engelm.) Small, Fl. S.E. U.S. 969. 1903. LCCTOTYPE (Yuncker 1921): U.S.A.
Engelmann Lindheimer!"
Western Louisiana, [no Gregg (MO). also cited "Texas,
datel,J. s.n.
The only between var calyptrata and var gronovii apparently are
differences
in corolla size and the way the corolla is shed. In var. calyptrata, as the capsule
expands, the corolla breaks transversely more or less regularly from the base
proximal portion
and permanently narrowed portion of the tube holds the of
a
the corolla (including the styles and remnants of the stamens) coherent as it is
breaks irregulady trans-
atop the capsule. In var gronovii, the corolla
lifted
expands and remains the cap-
versely and longitudinally, as the capsule it at
might be intermediate, the corolla tube
sule base. In the few plants studied that
narrowed and breaks irregulady but lifted at the top of the
not apically is
is
most narrowly
some Variety calyptrata the dis-
capsule at least in flowers. is
C
complex and perhaps the most sharply de-
tributed entity of the gronovii
C
cephahnthi unique
the
fiowever, var calyptrata very similar to in
fined, is
m
commonly 4-
way corolla caps the mature capsule, but the latter differs
and depressed-globose capsules.
merous narrower infrastaminal
flowers, scales,
The between these three taxa need further study.
relationships
Flowering Aug-Oct.
and ecology-V.S.A.: Louisiana, Texas.
Distribution
woody and herbaceous
Hosts: species.
imperiled imperiled) (T2,
Conservation status.-TlTl (critically to
NatureServe 2005).
204
BRIT.ORG/SIDA
22(1)
Cuscuta umbrosa Hook,
2. Beyr. ex Fl. Bor. Amer. 2:78. 1840. (Fig. 1 b, d, h).
g,
Grammica umhwsa WA.
(Beyr. ex Hook.) Webci; Southwest. Naturalist Lectq-
18:319. 1973.
TYPE (designated here): "Northwestern America;' Douglas Among
(K). three collections,
s.n.
Hooker C
cited ^'N.W. Coast of America, Douglas." The interpretation of umbrosa has been
mtertwnied with that of C.gyonovii var curia (below), apparently obscuring an unequivocal
name
lectotypification of the earlier from Hookers publication. Engelmann (1859) placed "C
umhroui Hook- synonymy
Ic. in part" in of C. uinhrosa var. curia, and he cited "Northwestern
America, Douglas! Fremontr Yuncker (1965) treated Cuscuta gronovii var lu rta as a synonym
of C. uDihrosa but noted only that the type locality of umbrosa was ^^North western America."
C.
Cuscuta gronovii Willd. ex Roem.
Schult. var. curia Hngclm., Trans. Acad. Louis
Osr Sci. St. 1:508.
1859. Cuscuta cii rta (Fngelm.) Rydb., Bull. Torrey Bot. Club 40:466. 1913. Lectotype (Yuncker
America
1921): '^Northwest (Fremont's 3rd Fxped, 79), the type, in the Engelmann lerb. (MO).
1
C C
For curia (with umhrosa as a synonym), Yuncker (1932) later cited "Northwestern
Kew
America, (Douglas, a specimen in the Herbarium)'" as the "type."
Cuscuta mcgahcarpa Rydb., Torrey Club LECTOTYPE
Bull. Bot. 28:501, 1901. (formally designated
here. Reveal in herb. 1977): U.S.A. Colorado: Cucharas Creek, near La Veta, 7000 15 Sep
ft,
Vredand
1900. 670 (NY). Rydbcrg also cited Tweedy 227S, a collection from Sheridan
Co.,
Wyoming.
medium
Stems mm)
yellow
to coarse, to orange. Inflorescence of pedicellate (1-7
commoiily
lowers cymose
in dense, paniculate each flower with one
clusters;
t
ovate broadly
to triangular bract at the pedicel base. Flowers 5-merous (some-
mm
times 4-merous), 2-3.5(-4.4) long
not papillose, white-cream, with few
a
,
many
round
to isolated, to ovoid translucent laticifers in the calyx, corolla and
ovary. Calyx campanulate reaching to ca. the middle of the corol tube, divided
la
ca. 1/2-2/3 the calyx length, lobes ovate, apex rounded or obtuse, basaUy over-
lapping, margins mm,
entire to serrulate; corolla tube campanulate, 17-2.3(-2.7)
lobes mostly ovate
to broadly-triangular ovate, rounded-obtuse, 1/3-1/4
the
tube corolla tube, spreading or ref lexed; epicuticular wax absent or reduced
to
few
a reticulated rodlets; stamens exserted, anthers on filaments equaling
or
longer than
anthers; pollen 3(-4)-zonocolpate, (18-)20-25(-28)
^im long,
rounded
or truncated polymorph same
at poles, in the anther or flower, from
spheroidal to subprolate (the former shape more frequent); usually tectum
perforatum
to almost microreticulate, puncta 0.4-0.7
|.im in diameter, rarely
tectum imperforatum;
sexine scabrate with
isolated granules; infrastaminal
scales infrastaminal and
scales broadly-oblong, apically truncate
to slightly
1/3) distinct, slender, thickened
mm
at the base 0.3-0.7(~0.9) long; stigmas capitate, globose. Capsules
subglobosc, ovoid to globose-conic or subobpyriform, thickened
the
at top,
oppening
interstylar relatively small, sometimes with short and broad but
a
x mm,
discernible neck, 3.5-6.5(-7) 3-5(-6) indehiscent or irregulat^y dehis-
cent. Seeds 3-4 per capsule. (1.8-)2-2.5(-2.8) X 1.5-1.65 mm, dorsoventrally com-
pressed to obscurely angled, broadly-elliptic obovate; hilum subrotund, 0.40-
to
mm
X mm,
0.60 0.39-0.46 vascular funiculum
scar of 0.20-0.25
linear, long.
COSTEA ET AL, TAXONOMY OF CU5CUTA GRONOVII AND UMBROSA 205
C.
vertical to slightly oblique. Surface of seed epidermis alveolate v^hen dry and
papillate, with cells 20-50 |im in diameter
Cuscuta umhrosa similar in some aspects of flower, and seed mor-
is fruit,
C
phology Both have infrastaminal than
to rostrata. scales shorter the corolla
and and
tube large capsules seeds. Cuscuta rostrata recognizable by the nar-
is
rower infrastaminal and by prominently beaked ovary and
scales especially the
capsule. Cuscuta umhrosa, however, sometimes produces capsules with a short
may C
and broad neck, which indicate homology with the beaked capsules of
2n-l
rostrata.
NW
—CANADA:
and
Distribution ecology Alberta., Manitoba, Ontario (only
one collection), Saskatchewan. U.S.A.: Colorado, Kansas, Minnesota, Montana,
New New
Nebraska, Mexico, York reported by Mitchell and Tucker
(as 1997;
North Oklahoma, Wyoming.
possibly a waif), Dakota, South Dakota, Utah, Flow-
ering Jul-Qct. Growing in shaded and/or wet environments. Hosts: species of
Salix, Ampelopsis, Symphoricarpos, Clematis, Epilohium, Convolvulus,
Linum, and
Scutellaria, other genera.
G4G5
Conservation status.— (apparently secure to secure) (G5, NatureServe
2005).
APPENDIX
L
VOUCHERS
FOR SEM STUDY
TliE
CANADA.
la. Cuscuta gronovii yar gronovii (16 collections examined). ONTARIO. Leeds Co.:
km
Rear of Leeds Twp., knn N of Oak Leaf, 6 5E of Delta, 44 35'20"N 76 02'55"N, 3 Sep 990,
1
1
NOVA
Catling 5111 (DAG). SCOTIA. Lunenburg Co.: Wentzell LakeU7Aug ]92]Jemald & Long
ALABAMA. Montgomery
24402 (NY). U.S.A. Co.: Alabama River Bottom5,4 mi N of Montgomery,
ARKANSAS.Hot
31 Oct 1943,/(peooeretG/.5.n. (NY). Springs Co.: Magnet Cove,470ft2 Oct 1938,
Hale 2062 (NY). FLORIDA.Taylor Co.: vicinity of Adams Beach, 6 Oct 961 Godfrey & Honk 61 565
1 1
,
(NY).GEORGIA.
Fulton Co.: Atlanta, 11 64 Clifton Rd,l Oct 1956, ei/rbanc^s.n. (NY).INDIANA. Fran-
klin Co.: bank of river 4 mi S of Laurel, 2 Oct 924, Deam 41026 (NY). MAINE. Knox Co.: seashore
1 1
Aug
(NY),MASSACHUSETTS.BarnstableCo.:26 Aug
Martinsville, 14 ]9S2,Fnesners.n. 1928,FernGW
er fx/ccaraeGrayane 48(7 (NY). MICHIGAN. Kent Co.: 7 mi N of Lowell, 28 Aug 1931, Yuncker &
a/.,
Noxubee
Welch 3792 (NY). MISSISSIPPL Co.: 3 mi of Macon, 22 Oct 960, McDaniel2276 (NY). MIS-
1
W
SOURI. Jefferson Co.: 6.5 mi of DeSoto, 38 07'30"N 90 4030" ,25 Aug ]99], Raven & Raven
NEW
27930 (NY). Saint Louis Co.: near Saint Louis, 878, Eggert (NY). JERSEY. Middlesex Co.:
s.n.
1
New
opposite Brunswick, Oct Mackenzie OHIO. Ottawa Ottawa
]9]8, (NY). Co.: Turtle Creek,
s.n.
Aug PENNSYLVANlA.York
National Wildlife Refuge,570ft.22 1996,H/7/ 25542 (NY). Co.: near McCall's
Ferry 9 Sep ]S93, Heller & Halbach /357 (NY). WISCONSIN. Waushara Co.: 2.5 mi SSW of Borth, 15
WO
Aug
]984, Bewick {NY).
b. Cuscuta gronovii var latiflora (12 collections examined). U.S.A. INDIANA. Daviess Co.:
1
Deam SW
1.5 mi SE of Newberry,21 Sep 932, 53259 (NY). Pasey Co.: 12 mi of Mt. Vernon, 20 Aug
1
Deam Deam
1922, 37712 (NY). Sullivan Co.: along Busseron Creek, 2 mi E of Sullivan, 3 Oct 1931,
5/395 (NY). IOWA. Wapello Co.: Cliffland,NWl/4 Sec. T-71N,R-13W, 30 Sep ]954, Davidson 4051
11,
MARYLAND,
(NY). (no county given): at mouth of Little Gunpowder River, 2 Sep ]902, Shall 289
NEW
MICHIGAN. Kalamazoo
(NY). Co.: Portage Twp., 20 Jul 936, Hanes s.n. (NY). JERSEY. Mercer
1
NEW
Co.: Duck Island, below Trenton, 21 Aug 1921, A4ac/(enz/e (NY). YORK. Nassau Co.: Long
s.n,
206
BRIT.ORG/SIDA
22(1)
IslandOysterBay,"]! Sep 1926, Ferguson 609/ (NY). Suffolk Co.: Long Island, Water Mill, 10 Aug 1925,
Wayne
Ferguson 4293 (NY). Co.: Lake Ontario, Fairbanks Point,WilliamsonTwp., 246 10 Aug 1939,
ft,
H/n/cey4484(NY).TEXAS.WoodCo.:LakeElli5,3Sep n750 VIRGINIA.
1942, ^ande// (NY). Prin
Anne
Co.: along North Landing River, near Creed's, 9 Sep ]93S, Fernold etal. 501 (NY).
1
1c. Cuscuta gronovii var. calyptrata collection examined) U.S.A. TEXAS. Rockwall Co.:
(1
NW
3.25 mi of Rockwall, 22 Oct 946, Cory 52529 (SMU).
1
CANADA. ALBERTA:
Cuscuta umbrosa Aug
(14 collections examined). Lethbridge, 30
2.
MANITOBA:
(DAG). Awcme Aug (DAG);
\97i. Alien 75(7 H), 9 (no year), Criddle s.n. Delta,
&
Aug
S13-T14-R7, 20 1953, Ldre Love 6199 (DAG); Delta Marsh, University Field Station,
50*^1 1'N 98" 24^W, 9 Aug 1984, Szumigalski 18 (DAO); Otterburne, 13 Aug 1954, Bernard 497
SASKATCHEWAN:
(DAG); Winnipeg, Mulvey Avenue, Sep 1975, F/eW^' (DAG). Carnduff,
a.//.
Aug
20 \94E, Biuh/& Lodge 7/97 (DAG); 10 mi S of- and 0.5-1 mi E of Gainsborough, Hutlcr
COLO-
River Valley, NHl/4 Sec. 9 Tl R30 Wl, 29 Jul 1988, Harms 39594 (DAO). U.S.A.
RADO,
Aug MIN-
Teller Co. (?).: Manitou (Experimental Forest'.^), 3 1892, Mulfords.n. (NY).
1
SW
NESOTA. &
Carlton Co.: 3 mi of Fond du Lac, 22 Aug 1945, Moore Hujf 18199 (DAO).
NORTH
DAKOTA. SOUTH DAKOTA.
Cass Co.: Fargo, 9 Sep 1950, Stevens 1228 (DAO).
& UTAH.
Potter Co,: Forest City, Sep 1892, Glosser (NY). Salt Lake Co.: City
Griffiths s.n.
Aug
Creek Canyon, 8000 10 1883, Leonard 250 (NY).
ft,
ACKNOWLEDGMENTS
Dan Musselman and an anonymous
Austin, Lytton reviewer provided valuable
comments
and suggestions that improved an earlier version of the manuscript.
We
thank directors and/or curators from ACAD, ALTA, ARIZ, ASU, BRIT, DAO,
MEXU
HAM, MTMG,
GH, MICH, MT, NFLD, N5PM, OAC, QFA, QUE, RBG,
F,
& &
TEX UC UN U WO,
RSA, SASK, SFS, LL, TUP, UBC, JEPS, UNB, M, US, USAS,
UWPG, WTU, XAL
WAT, WIN, WIS, and
for loans to Costea. Special apprecia-
NY
tion extended to staff for approving and preparing the numerous volumi-
is
nous loans. Robert Vogt and Meine Jutta (B) provided helpful information
re-
4
garding the type coflection of Cuscuta gronovii, and Kaj VoUesen (K) provided
information regarding the type collection of Cuscuta umhwsa. Alexandra Smith
assisted us with the scanning electron microscope and Elma Sch checked
w^eigert
the translation of the Spanish abstract.
REFERENCES
Costea, M., G1. Nesom, and Stefanovic 2006. Taxonomy of the Cuscuta pentagona com-
S.
plex (subsect./lri/erj5e5:Convolvulaceae) North America. Sida 22:151-175.
in
and The dodders
CrinsAVJ. B.A.Ford. 1988. parasitic (Cuscuta: Cuscutaceae) Ontario.
in
CanadTield-Nat. 209-21
02:
5.
1
Darbyshire, 2003. Inventory of Canadian agricultural weeds. Agriculture and Agri-Food
S.J.
Canada, Research Branch,Ottawa, ON. 396
pp.ElectronicPublication;<http://res2.agr.ca/
ecorc/weeds_herbes/title-titre_ehtm>.
A
Dawson, H. 1984. vegetative character that separates species of Cuscura.Proc. 3rd Internt
J.
Symposium on Weeds. 84-1
Parasitic Pp.1 87.