TAXONOMY OF THE CUSCUTA INDECORA (CONVOLVULACEAE) COMPLEX IN NORTH AMERICA

2006·12.1 MB·English

by  Mihai Costea

TAXONOMY
OF THE CUSCUTA INDECORA
(CONVOLVULACEAE) COMPLEX NORTH
AMERICA
IN
Mihai Costea Guy Nesom
(corresponding author)
L.
Department
of Biology Botanical Research ofTexas
Institute
Wilfrid Laurier University 509 Pecan
Street
75 University Avenue West Fort Worth^ Texas 76102-4060,
U.S.A.
Waterloo, Ontario
[email protected]
N2L3C5XANADA
[email protected]
Sasa Stefanovic
Department
of Biology
University of Toronto at Mississauga
3359 Mississauga Road
Mississauga, Ontario
L5L1C6XANADA
ABSTRACT
m
The Cuscuta indecora complex {Cuscuta subsect. Indecorae) North America characterized by
is
fleshy, papillose flowers, corolla lobes with mflexed tips, and capsules with a thickened stylopo-
C
dium. It includes Cindecora, coryli, C. warneri, and C.jepsonii- Recent taxonoinists have proposed
C
We C
to treat the latter three taxaat varietal rank within inderora or as synonyms of maintain
it.
warneri and C.jepsonii at specific rank because of their distinctive morphology, even though the
known
latter is only from the type collection and the former from a very limited number of speci-
mens. Cuscuta indecora includes var. indecora, var. hngisepala, and C. indecora var. aticnuata (Wa-
comb, nov
terfall) Costea, et stat. Cuscuta indecora vars. bifida and neuropetala are treated as syn-
onyms C
of CAndecora var ndecora. Cuscuta warneri and coryli are reported as new floristic records
i
New
from Mexico and Quebec,
respectively.
RESUMEN
El complejo Cuscuta indecora (Cuscuta subsect. Indecorae) en America del Norte esta caracterizado
por flores papllosas y suculentas, lobulos de la corola con puntas inflexionadas, y capsulas con un
estilopodio engrosado. El complejo incluye C. indecora, C. coryli, C. warneri. y C.jepsonii.
Recientemente algunos taxonomos han propuesto cratar las ultimas tres variedades dentro de la
C
C
como
categoria indecora o un sinonimo del mismo. Mantenemos que warneri C.jepsonii estan
y
en una
categoria especial por su morfologia distinta, a pesar de que la ultima se conoce solo por la
coleccion primera conocida mediante un numero muy
del tipo, y la es de especimenes limitado.
Cuscuta indecora incluye var. indecora, var. longisepa/a, y C. indecora var. attenuaia (Waterfall) Costea,
C
comb, et stat. nov Cuscuta indecora vars. bifida y neuropetala son tratadas como sinonimos de
indecora var. indecora. Cuscuta warneri y C. coryli figuran como nuevas citas floristicas de Nuevo
Mexico Quebec, respectivamente.
y
Cuscuta subsect. Indecorae Yuncker characterized by fleshy, papillose flow-
is
corolla lobes with inflexed and capsules with thickened stylopodium
ers, tips, a
SIDA22{1): 209-225. 2006
BRIT.ORG/SIDA
22(1)
210
North American
Yuncker included three spe-
(Yuncker 1932, 1965). In view,
s it
C
C
Engelm, and warneri Yuncker) and one
(C
indecora Choisy, coryli
cies
C
A Wa-
from South America (C sienolepis Engelm.). fifth species, attenuata
akhough
and
was added subsect. Indccorae by Prathcr Tyrl (1995),
to
terfall,
C C
Waterfall (1971) had suggested that attenuata resembled compacta]nss. of
CAndecora,
Lepidanche Engelm. Cuscuta attenuata closely similar to
subsect. is
emphasizing
and Tyd maintained both rank, their
but Prather (1995) at specific
was
apparent reproductive Cuscuta warneri treated by Beliz in her
isolation.
C
unpubhshed combination
Ph.D. thesis (1986) as a variety of indecora; this
some North American overviews Kartesz
has been included recent 1999).
in (e.g.,
Cuscuta jepsonii Yuncker, which was initially treated in subsect. Calijornicae
C
with
Yuncker (Yuncker has been recently considered conspecific
1932),
indecora 2002). The taxonomic status and relationships of these
(Beliz 1993,
taxa are reevaluated here, based on morphology and micromorphology of flow-
and
capsules, seeds, pollen.
ers,
METHODS
morphology based on samples from specimens from NY,
Descriptions of are
which herbarium (Appendix Measurements and pictures
includes Yuncker's
1).
KY
Samples
were taken with a scanning electron microscope Hitachi S-570 at 15
nm
K
were coated with 30 gold using an Emitech 550 sputter coater. Terminol-
micromorphology and and pollen
ogy regarding the of flowers, seeds capsules,
m
were described in detail in the first paper published this issue (Costea et al.
2006). Conservation status was assessed using NatureServe (2005) ranks and
criteria.
TAXONOMY
—
and
Cuscuta indecora and C. coryli. Cuscuta indecora a highly variable
is
common species both in North America and South America (Engelmann 1859;
&
Yuncker 1921, 1932. 1965; Beliz 1986; Prather Tyrl 1995). Cuscuta coryli is less
C
and sympatric with indecora over significant geographic area
variable a
is
C C
below). Yuncker (1932) mentioned that coryli closely related to
(see "is
commonly
which
indecora, but distinguished by often 4-parted flowers are
is its
mm
We
and
smaller 2 long), rudimentary scales shape of fruit." find that
(ca.
C
flowers of coryli may be both 4-merous and 5-merous on the same specimen,
and sometimes even 3-merous. Engelmann (1843) noted "flowers frequently 5-
C
parted" for coryli, but he later (1859) modified this observation to flowers
many
Although and appar-
"mostly 4-parted." the type collections of C. coryli
ently have more 4-merous than 5-merous flowers, 5-merous flowers predomi-
nate in some specimens. Ranges of variation in floral size in C. indecora 3.nd C.
mm
Cuscuta indecora has small flowers, 2.1-2.7 long,
coryli are closely similar.
JAXONOMY
COSTEA ET AL. OF CUSCUTA INDECORA COMPLEX 211
mm mm
C
with the coroha tube 0.9-L5 long; flowers of coryli are L7-2.6(-3)
mm
long, with the corolla tube (0.7-)l-L4 long.
GH)
Such observations perhaps led Beliz (1987-1988, in herb. NY, to con-
C
sider Cuscuta coryli conspecific with indecora. Although they are clearly re-
lated and sometimes overlapping in several character states, the two taxa can
be usually distinguished using a combination of characters (see key and de-
and
scriptions below) their recognition as separate species appropriate.
is
Description of infrastaminal scales in these two species has been confus-
m
Engelmann was
ing. For example, Cuscuta coryli, consistent, describing the
infrastaminal scales as "appressed, bifid, consisting of a few teeth one or two
...
teeth on each side of the filament (1942) or lobes laterally adhering to the
...
lower (attached) part of the filament" (1859). Yuncker (1921, 1932) added "scales
rudimentary, bifid, toothed, ordinarily reduced to toothed wings on either side
filament attachment." Figure 42e depicted one infrastaminal
of (1921) bifid scale
with 1 or 2 fimbria on the each side of the staminal filament, while in Fig. 39
(1932), these lateral fimbria are replaced by dentate lobes (wings). In his 1965
treatment, Yuncker removed the term "bifid" from the description of
C
infrastaminal scales of coryli 'dnd applied to C.indecora var bifida Yuncker,
it
which was
described as identical in other respects to C. indecora var indecora.
We
find that the infrastaminal scales of var bifida are not truly bifid, but rather
may
which
the spathulate scales have 2 or 3(-4) deeper apical incisions, create
may
same
2 or 3(-4) lobes that are further fringed. "Normal" scales occur in the
flower together with lobed ones. Such plants are regarded here as populational
Engelmann
variants of C.indecora van indecora. Bifid scales, as pointed out by
C
(1842, 1859), are characteristic of coryli.
—
The and Yuncker
varieties of C. indecora the status of Cuscuta attenuata.
(1965) treated Cuscuta indecora with three varieties (var. indecora, var. bifida,
and van noted we maintain var
longisepala): as below, tentatively longisepala
but var bi/idaisnotappropriatelyrecognized. Cuscuta var neuropetala
zncieco ra
(Engelm.) Hitchc, distinguished by relatively larger flowers, has been ac-
its
We
cepted in recent overviews of the genus (Beliz 1993, 2002; Kartesz 1999).
find that although var neuropetala can be often identified, connection to var
its
indecora by a series of intermediates makes taxonomic recognition unfeasible.
and
Prather Tyrl (1995) observed that Cuscuta attenuata morphologically
is
C
similar to indecora. Indeed, Yuncker annotated (in herb. NY) some collec-
C
The
tions of attenuata as C.indecora var longisepala. present study substan-
the w^hich extend the morphology and micromorphology
tiates similarities, to
C
and The
of perianth, seeds, pollen, capsules. geographic range of attenuata
completely within that of var indecora, but preliminary evidence suggests
lies
C UPGMA
may
and
that attenuata have narrower host preference. In princi-
a
C
pal components analyses by Prather and Tyrl, attenuata part of a single
is
BRIT.ORG/SI0A
212 22(1)
C
cluster with indecora var. indccora and var. longisepala; univariate analyses
separated these three taxa, although they were broadly overlapping in morphol-
ogy. Prather and Tyrl 456) concluded that "C attenuata a distinct species
is
(p.
C
albeit morphologically similar to indecora. In the absence of reproductive
isolation we might treat C.attenuala asa variety of C. indecora." Experimental
crosses by Prather and Tyrl between attenuata and both varieties of
C. C.
indecora produced neither fruits nor seeds, while populations of C. attenuata
were Whether indecora and var longiscpala were mterfertile
interfertile. var.
was
not reported.
For consistency with the degree of morphological difference between other
C
we
Cuscuta species (as are recognizing them), attenuata treated here at
is
C
varietal rank within inciecora.ln view of apparent reproductive isolation
its
more
and host specialization, a strongly defined entity than var hngisepala,
is
it
which also broadly sympatric with var indecora. Var longiscpala istentatively
is
may
maintained here, until biology and evolution be better understood.
its
Cuscuia wanieri.— This species a strikingly distinct dodder Each calyx
is
lobe apically prolonged into a conical spur-like projection (Fig. 4 a,b,d),
is
and and
infrastaminal scales are oblong with truncate dentate apex, capsules
have a collar-like apex and very short styles (Fig. 4d). The calyx spurs are
m
maximum
when
and
accrescent, relatively small flower reaching size in fruit,
they detach easily (at least on dry material). Cuscuta warneri shows strong simi-
morphology and micromorphology
larity to C. indecora in of perianth, cap-
sules, seeds, and pollen (as also observed by Yuncker 1960). Beliz (1986) wTOte
C
abnormal
warneri probably an
that "critical studies, how^ever, indicate that
is
C C
specimen and more
of indecora until material available for studies,
is
...
C
Our
warneri recognized as a variety of indecora.'' observations, in contrast,
is
C
do not indicate that warneri teratological. Flowers have all the components
is
and they are each capsule usually with 2 seeds in which the embryos
fertile,
C
known
appear develop normally. The that warneri only from the
to Fact
is
New
type locality and a collection from Mexico (see bellow) indicates that its
distribution localized, but morphological distinction justifies continuing
is its
recognition specific rank.
at
The spur-like projections of the perianth of Cuscuta warneri have an un-
knowm morphology encountered more
biological Similar also in other,
role. is
C C
distantly related species: runyonii Yuncker, appJanata Engelm.,
C.
C
Urban, and chapalana Yuncker
holdinghii
—
Cuscuia jepsonii. This species was described and included by Yuncker
(1921, 1933) in subsect. Calijornicae because infrastaminal scales are absent or
synonym
reduced to ridges. Beliz (1986) initially considered to be a of C.
it
&
califor nica Hook. Arn. var. papiUosa Yuncker, but she later (1993) treated
it
C
synonym
as a of indecora var. indecora. She did not provide substantiating
UC
knowm
evidence for these decisions. The species only from the holotype.
is
COSTEAETALJAXONOMYOFCUSCUTAINDECORA COMPLEX 213
may
Based on study we confirm belong
of this collection, that C.jcpsonii to
subsect. Indecorae, where represents an extreme case of infrastaminal scale
it
C
reduction. The 5-merous flowers and papillae morphology are similar to
mm
indecora, from which differs by very small anthers, 0.2-0.3 long, which
it
C
more
are like those of coryli.
many
some
Infrastaminal scales vary to extent in Cuscuta species. Never-
know
theless, we of no instance of complete reduction of infrastaminal scales
where development normal, even
in a species scale characteristically vari-
is if
For this reason, and until additional material can be studied, C.jepsonii
able. is
maintained
as a distinct species.
KEY TO THE OF CUSCUTA INDECORAE NORTH AMERICA
SPECIES SUBSECT. IN
Infrastaminal scales absent or reduced to ridges 4.C.JGpsonii
1.
Infrastaminal scales present.
1.
Calyx lobes with an apical spur-like projection; infrastaminal scales dentate at
2.
apex 3.C.warneri
2 Calyx lobes without spur-like projections; infrastaminal scales fimbriate.
mm
commonly
Flowers 5-merous, 2-5.3 long Jnfrastaminal scales united with
3.
the corolla tube for 1/3-1/2 (rarely 3/4) of their length, subspathulate to
spathulate, rarely 2-3 lobed,apex rounded rarely truncate, with (6-)20-35(-
± when
50) fimbria; capsule yellowish and semi-transparent dried, globose,
subglobose to slightly depressed-globose, 0.8-1 times wider than long, the
.5
suture lines between the 2 carpels not or only slightly depressed C. indecora
1
.
mm
Flowers (3-)4-5-merous, .7-2.6(-3) long;infrastaminal scales united with
3.
1
the corolla tube for most of their length, oblong, bifid with 1-3 fimbria on
±
3-6
each side of filament attachment, rarely truncate with fimbria;capsule
brown and not semi-transparent when dried, globose, becomes
initially
between
depressed, 1.6-2.4 times wider than long, the suture lines the 2
carpels depressed, forming longitudinal groove on the opposed sides of
a
capsule 2. C. coryli
Mem.
Geneve 1842
Cuscuta indecora Choisy, Soc. Phys. 9:278, t3, (Fig.
1, f.5. 1,
Fig. 3a,d,e,g,h). Cuscuta decora Choisy ex Engelm. indecora (Choisy) Engelm., Trans.
var.
Grammica
Acad. Sci. St. Louis 1:502. 1859. indecora (Choisy) W.A. Weber, Southw. Naturalist
&
Amer
Epithymum indecorum (Choisy) Nieuwl. LunelK Midi. Naturalist
18:319. 1973. 4:511.
1916. Type: MEXICO. [Tamaulipas]: "Mexicum ad Matamoros," Berlandier 2285-865 (holo-
G-DC; 1S0TYPE5: MO,
TYPE:
P).
mm
pan-
Stems 0.4-0.7 yellow orange. Inflorescences loose dense,
thick, to to
iculate-cymose sometimes originating endogenously; pedicels 0.5-6
clusters,
mm
long, papillate-hispid to glabrous; bracts one (rarely 0) at the pedicel base,
mm
ovate lanceolate. Flowers (4-)5-merou5, 2~5.3 long fleshy translucent-
to
when when
white same dark-brownish epidermal
fresh, of the color or dried;
arranged in rows, fleshy with anticlinal walls convex (dome-like), addi-
cells
when
tionally with cylindric-conical papillae 40-80 |am long; flowers dry up,
rows of dehydrated epidermal cells are usually easily discernible; epicuticular
wax
represented by longitudinally reticulated rodlets; laticifers isolated or in
214 BRIT.ORG/SI0A
22(1)
=
FiG.l. Floral variation in Cind€cora:a~e, var.indecora (scale bar mm);f. Papillae on the calyx of var.indecora (scale
1
=
mm).
bar 75 jam); g. var, longisepala; h, var. attenuata (scale bar
1
COSTEA ET TAXONOMY OF CUSCUTA INDECORA COMPLEX 215
AL.,
longitudinal rows, ovoidal to elongated, present in the perianth along the
midveins, ovary and capsules often running longitudinally and yellowish-or-
somewhat
ange colored. Calyx cupulate, 1/2 to longer than the corolla tube, di-
vided 1/2-2/3 of the length, lobes triangular-ovate to lanceolate, acute to at-
more
tenuate, or less overlapping at the base. Corolla tube campanulate,
mm
campanulate-cylindric, subglobose or suburceolate, 1-3.2 long, lobes 0.7-
mm
1.5 long triangular-ovate, acute, ca. 1/3 to equaling corolla length, suberect
Siamens
to erect, apically inflexed. barely exserted or enclosed; anthers elliptic
mm
x
to oblong, (0.5-)0.6-0.9(-1.2) 0.3-0.5 long; filaments equaling or longer
24-36
than anthers; pollen 3(-4)-zonocolpate, subprolate to prolate, jam long,
tectum impcrforatum or with a few isolate puncta, sexine scabrate with iso-
lated granules. Infrastaminal scales reaching the filaments, united with the
corolla tube ior 1/3-1/2 (rarely 3/4) of their length; subspathulate to spatulate,
apex rounded, rarely truncate or 2-3(-4) lobed, with (6-)20-35(-50) fimbria.
mm
weakly
Styles distinct, 1-2.5 long, ± unequal, evenly filiform, suberect or
divergent; stigmas capitate, globose. Capsules yellowish to light-brown and ±
when
semi-transparent dried (pericarp glabrous, subglobose, globose,
thin), to
slightly depressed-globose, 0.9-1.5 wider than long, narrowed and thickened
around the style bases, indchiscent or irregularly dehiscent, surrounded or
2-4
capped by the withered pericarp epidermis smooth. Seeds per cap-
corolla;
sule, 1.42-1.86 X 1.25-1.6 mm, shape heterogeneous on the same plant: dorsoven-
trally compressed to weakly angled, broadly elliptic to transversely oblique,
x
hilum subtermmal, rarely almost terminal, broadly 0.40-0.45 0.32-
elliptic,
mm; mm,
0.36 vascular scar linear, 0.15-0.18 vertical; seed surface variable: a)
when
epidermis cells more or less polygonal and puzzle-like, b) alveolate dry
when
and papillose wet, and c) only some cells are papillose and the rest are ±
20-50
epidermal
puzzle-like; size of cells |.im in diameter.
KEY TO THE OF CUSCUTA INDECORA
VARIETIES
Calyx lobes ovate triangular, reaching ca. 1/2 of the corolla tube a.C. indecora
1.
indecora
var.
Calyx lobes lanceolate mostly longer than the corolla tube.
1
.
Z Calyx lobes acute; flowers in loose clusters; parasitic on a wide range of species
indecora longisepala
b. C. var
2 dense on annua
Calyx lobes attenuate;flowers in clusters; parasitic primarily Iva
indecora attenuata
c. C. var.
Cuscuta indecora var indecora. neuwpctalaEngdm., AmcL
a. (Fig. 1 a,b,c,d,e). CusciUcJ
Sci. Arts 45:75. 1843, Cuscuta indecora Choisy var neuropetala (F^.ngclm.) Hitchc, Contr.
J.
U.S. Natl. Herb. 3:549. 1896. TYPE: U.S.A. TEXAS. [Harris Co.]: in wet prairies near Houston, on
different Compositae,such asLialris, Solidago, Hclianlhus, Rudbeckia/aud on Myrica ccrifcra,
IM
Aug Lindheimer MO;
1843, (holotype; ISOTYPES: NY, US).
Cuscuta ncuropelala Engelm. var liitoralis Engelm., Boston Nat. Hist. 5:223. 1845. Lectotype
J.
(here designated; Beliz 1986, in herb.): U.S.A. TEXAS. [Galveston Co.]: Gaiveston, Apr 1843,
216 BRIT.ORG/SIDA
22(1)
Lindhcimersn- (MO; The
Isolectotypi:: NY). protologuc noted Galveston
'^Scashore of Island,
on lycjum caroliniamim, Bonichiajrutcsccus, Ivajrutcscens, etc, Flowers in May."
Cuscida vc/'niarsc?P:ngcbn. var. Iijs/;icinlcj Fngelm, Amer. Sci, Arts 43:341. 1842. Cuscutd indccora
J.
var. hi^ndula (EngclmJ Yunckcr, llMnois Biol. Monogr. 6:148, 1021. TYPE: U.S.A. TEXAS.lHarris
^1n dry and sterile prairies, of louston, on Eulhiimia, Sthraukia, Asia\ Amhrosia,
Co-l: \\'est
1
EvoJyiilu^, and other low herbs, flowering in April and May. Lindhcimcf' (HOI.OTYPE:
s.n.
J\
MO, 2285-MO
presumably). Yunekcr (192]) cited Bcrlandier the type of indecora
C. var.
i\s
h ispidiild.hui that collection interpreted here as the holotype of ndccora indccora).
is C. (var.
i
Amer
Cuscuta hlspidulu tZngelin., Sci. Arts 45:75. 1843. TYPE: U.S.A. TEXAS. larris Engel-
Co.];
J. [1
mann
did not cite a collector but noted "in dry and sterile prairies west of Houston. Flower-
ing in April and May;' apparently referring to the same Lindhcimcr collection typifying
C.
verrucosa var hlspidula (above). Cuscuta hispidula apparently was not intended by Fngeb
mann
new
to be a combination based on the eadier C. verrucosa Fngehn. var. hispidula, as he
C
wrote ''Cuscuia hispidu la n. sp." and "Compare the remarks nrade in Vol. XLIII 34h under
p.
verrucosa.''
Cuscuta indccora var. bifida Yuncker, IHinois Biol. Monogr 6:14^J. 1921. Lectotype (designated
UC
here): U.S.A. NEVADA. Twin Springs. May-Qct. Pu rpus 6343 (UC 124538). Two sheets are
at
marked as "isotype"oi var. bifida: the other is Furpuss.n.. wnthout date (UC 124541).
CANADA:
and
Distrihution ecology— Saskatchewan. Alabama, Arkan-
U.S.A.:
sas, Arizona, California, Colorado, Connecticut, Florida, Georgia, Idaho, Iowa,
Kansas, Kentucky, Louisiana, Maryland, Michigan, Minnesota,
Illinois, Missis-
New
sippi, Missouri, Montana, Nebraska, Nevada, Neu^ Mexico, North
Jersey,
North Oklahoma,
Carolina, Dakota, South Carolina, South Dakota, Tennessee,
WEST
Washmgton, MEXICO;
Texas, Utah, Virginia, West Virginia, V\/yoming.
SOUTH
AMERICA. common
INDIES; probably most dodder
third
It is tlie in
North America, and
after C.gronovii C. avnpestris. hsted as a "locally" im-
It is
&
weed
portant in the U.S.A. and Argentina (Parker Riches 1993). Flowering
Jul-Nov. Hosts: wide range of herbaceous and woody Agalmis,
species,
e.g.,
Bciccharis, Borrichuh Chenopodium, Eupatorium, Hdianihus, Uctcwthcca, Hy-
pericum, Ipomoca, Iva, Kostclctzkyn^ Lcpidhim, Ligustrum, Myrica, PJuchca,
Polygcnnim,Rhynchosic\,Solidcigo,Suacdci,Symphyotrichum^Tcphro5ici,Vcr^^^^
G5 (common)
Conservation status— (NatureServe 2005). n - 15 (Pinkava
et
&
In = 30 (Pazy Plitmann
1974); 1995).
al.
Cuscuta indecora longisepala Yuncker, Monogr.
b. var. Illinois Biol. 6;149, Fig.
44, 97. 1921 (Fig. Ig). Typh: U.S.A. Texas. On the Blanco, Wright (llOLOTYPH; MO),
soi.
Distnhutio}] and MEXICO; SOLJTH AMERICA.
ecology —\J,S, Texas. Flower-
A.:
ing summer-fall. Hosts: herbaceous and woody Conservation
species.
status:
T2T1
(imperiled to critically imperiled) (not yet assessed by NatureServe 2005).
Cuscuta indecora comb.
c. var. atteiiuata (Waterfall) Costea, &r stat. nov. (Fig.
111). Cuscuta attcnuata Watcrlall, Rhodora Oklahoma. McCurtani
73:575. 197]. TVPI:: U.S.A.
Co.: Waterfall Creek, 8 mi S and 2 mi E of idabel, Waierfall 17157 (llOLOTYPE: OKLA, ISOTYPE:
GH).
&
Distribution and Oklahoma,
ecology —\J,S.A,: Kansas, Texas (Prathcr Tyrl
1993).
COSTEA ET AL, TAXONOMY OF CUSCUTA INDECORA COMPLEX 217
Conservation status.— imperiled (G2) (Natureserve 2004). Flowering late
Aug-Oct. Hosts: Iva annua, rarely Symphyotrichum spp., mudflats, floodplains,
and
disturbed
areas.
T2T1
Conservation status.— (imperiled to critically imperiled) (G2,
&
Natureserve 2005). 2n = 30 (Prather Tyrl 1993).
2. Cuscuta coryli Engelm, Amer. Sci. Arts 43:337. 1842. (Fig. 2, Fig. 3 b, c, f).
J.
Type: U.S.A. MISSOURI. [St. Louis Co]: "on Corylus near St. Louis,'^ Sep 1841, Engelmann s.n.
(hoi.OTYPH: MO; ISOTYPE: GH). The protologuc reads "on Corylus, in the barrens west of
St.
August and September. Tt
Louis, in
Amer
Cuscuta coryh Engelm. var. stylosa Lngelm., Sci. 43:337. 1842. TYPE: U.S.A. MISSOURI. iSt.
J.
Engelmann MO; GH,
Louis 'on Sohdago. Louis;' Sep 1841, lOLOTYPE: ISOTYPES: US).
Co-1: St. s.n. (l
The protologue reads "On Solidago, in dry prairies near Louis."
St.
Cuscuta inflexa Engelm., Trans. Acad. Louis L502. 1859 [nom. invahd.]. Apparently a renaming
St.
of Cuscuta coryli, the latter was cited immediately and first in synonymy (other names at
'ds
A
specific rank also were cited). number of collections were cited in the protologue.
mm
Stems 0.30-0.50 yellow orange. Inflorescences usually dense (some-
thick, to
times paniculate-cymose sometimes originating endogenously;
loose), clusters,
mm
pedicels 0.5-3 long, glabrous; bracts one at the pedicel base, ovate to lan-
mm
ceolate. Flowers 4-5-merous (rarely 3-merou5), 1.7-2. 6(-3) long, fleshy
when commonly
white dark-brownish w^hen epidermal fleshy
fresh, dried; cells
organized rows with anticlmal walls convex (dome-like); papillae those
in like
C
when
dry rows dehy-
described in indecora usually absent; flowers up, of
wax
more
drated epidermal cells are or less inconspicuous; epicuticular present
m
by
represented longitudinally reticulated rodlets; laticifers isolated or longi-
tudinal rows, ovoidal to elongated, present in the perianth midveins, ovary and
somewhat
capsules. Calyx cupulate, equaling or longer than corolla tube, rarely
in some flowers shorter than corolla tube, divided 1/2-2/3 of the length, lobes
triangular-ovate, acute, not or only slightly overlapping at the base. Corolla tube
mm
campanulate to suburceolate, 0.5-1. 3(-1.5) miTi long, lobes 0.8-1.2(~1.5) long,
triangular-ovate, acute, ca. 1/3 to equaling corolla length, suberect to erect,
apically inflexed. Stamens barely exserted or enclosed; anthers (0.2-)0.3-0.45
mm
C
X
0.19-0.25 long; filaments equaling or longer than anthers; pollen as in
indecora. Infrastaminal scales reaching the filaments, united with the co-
ca.
rolla tube for most of their length, oblong, bifid, with short dentate wings or 1-3
fimbria on each filament attachment, rarely trtmcate with 3-6 fimbria.
side of
mm
Styles distinct, 0.7-1.5 long, ± unequal, evenly filiform, strongly divergent
dark-brown and
in capsule; stigmas capitate, globose. Capsules not semi-trans-
when become
parent dried (pericarp thick), glabrous, initially globose later
evidently depressed, 1.6-2.4 times wider than long; the former suture lines be-
tween the 2 carpels forming a longitudinal groove on opposed sides of capsule;
depressed and thickened around the style bases, indehiscent or irregularly de-
surrounded capped by withered pericarp epidermis with
hiscent, or the corolla;
218 BRILORG/SIDA
22(1)
C =
Fig. 2. Floral variation in coryli: a-g. excepting e (scale bar mm); e. Convex fleshy epidermis cells in the corolla
1
=
(scale bar 150 |„im).